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"S.fijj! 


Journal  of 

Hymenoptera 
Research 


Volume  16,  Number  2 N^     /  /RPMafcS^ October  2007 

ISSN  #1070-9428 
CONTENTS 

GESS,  F.  W.  The  genus  Quartinia  Ed.  Andre,  1884  (Hymenoptera:  Vespidae:  Masarinae)  in 

southern  Africa.  Part  I.  Descriptions  of  new  species  with  complete  venation 211 

GRISSELL,  E.  E.  Torymidae  (Hymenoptera:  Chalcidoidea)  associated  with  bees  (Apoidea), 

with  a  list  of  chalcidoid  bee  parasitoids 234 

NEFF,  J.  L.  and  A.  W.  HOOK.  Multivoltinism  and  usage  of  multiple  nest  substrates  in  a  west 
Texas  sand  dune  population  of  Psendomasaris  phaceliae  Rohwer  (Hymenoptera:  Vespi- 
dae: Masarinae) 266 

PACKER,  L.  Mydrosoma  micheneri  Packer,  new  species,  a  new  diphaglossine  bee  from  Brazil 

(Hymenoptera:  Colletidae) 277 

PACKER,  L.,  A.-I.  D.  GRAVEL,  and  G.  LEBUHN.  Phenology  and  social  organization  of  Halictus 

(Seladonia)  tripartitus  (Hymenoptera:  Halictidae) 281 

PULAWSKI,  W.  J.  The  status  of  Liris  magnificus  Kohl,  1884,  and  Trachogorytes  costaricae 

R.  Bohart,  2000  (Hymenoptera:  Crabronidae:  Crabroninae,  Bembicinae) 293 

PUNZO,  F.  Interspecific  variation  in  hunting  behavior  of  Pepsis  grossa  (Fabricius)  and  Pepsis 

thisbe  Lucas  (Hymenoptera:  Pompilidae):  a  field  study 297 

SHIMIZU,  A.  and  R.  WAHIS.  Systematic  studies  on  the  Pompilidae  occurring  in  Japan:  genus 

Irenangelus  Schulz  (Hymenoptera:  Pompilidae:  Ceropalinae) 311 

WENG,  J.  L.  and  BARRANTES,  G.  Natural  history  and  larval  behavior  of  the  parasitoid  Zaty- 

pota  petronae  (Hymenoptera:  Ichneumonidae) 326 


INTERNATIONAL  SOCIETY  OF  HYMENOPTERISTS 

Organized  1982;  Incorporated  1991 


OFFICERS  FOR  2007 

Michael  E.  Schauff,  President 

James  Woolley,  President-Elect 

Michael  W.  Gates,  Secretary 

Justin  O.  Schmidt,  Treasurer 

Gavin  R.  Broad,  Editor 

Subject  Editors 
Symphyta  and  Parasitica  Aculeata 

Biology:  Mark  Shaw  Biology:  Jack  Neff 

Systematics:  Andrew  Deans  Systematics:  Wojciech  Pulawski 

All  correspondence  concerning  Society  business  should  be  mailed  to  the  appropriate  officer  at  the 
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MD  20705,  USA;  Secretary,  Southwestern  Biological  Institute,  1961  W.  Brichta  Dr.,  Tucson,  AZ  85745, 
USA;  Treasurer,  PO  Box  37012,  c/o  Smithsonian  Institution,  MNMH,  MRC168,  Washington,  DC 
20013-7012,  USA;  Editor,  Dept.  of  Entomology,  The  Natural  History  Museum,  Cromwell  Road,  Lon- 
don SW7  5BD,  UK. 

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to  The  International  Society  of  Hymenopterists.  Requests  for  membership  should  be  sent  to  the  Trea- 
surer (address  above).  Information  on  membership  and  other  details  of  the  Society  may  be  found  on 
the  World  Wide  Web  at  http://hymenoptera.tamn.edu/ish/. 

Journal.  The  Journal  of  Hymenoptera  Research  is  published  twice  a  year  by  the  International  Society  of 
Hymenopterists,  %  Department  of  Entomology,  Smithsonian  Institution,  Washington,  D.C.  20560- 
0168,  U.S.A.  Members  in  good  standing  receive  the  Journal.  Nonmember  subscriptions  are  $60.00 
(U.S.  currency)  per  year. 

The  Society  does  not  exchange  its  publications  for  those  of  other  societies. 

Please  see  inside  back  cover  of  this  issue  for  information  regarding 
preparation  of  manuscripts. 


Statement  of  Ownership 

Title  of  Publication:  Journal  of  Hymenoptera  Research. 

Frequency  of  Issue:  Twice  a  year. 

Location  of  Office  of  Publication,  Business  Office  of  Publisher  and  Owner:  International  Society  of 

Hymenopterists,  0  Department  of  Entomology,  Smithsonian  Institution,  10th  and  Constitution 

NW,  Washington,  D.C.  20560-0168,  U.S.A. 
Editor:  Gavin  R.  Broad,  Department  of  Entomology,  The  Natural  History  Museum,  Cromwell  Road, 

London  SW7  5BD,  UK. 
Managing  Editor  and  Known  Bondholders  or  other  Security  Holders:  none. 


This  issue  was  mailed  24  September  2007 


J.  HYM.  RES. 

Vol.  16(2),  2007,  pp.  211-233 

The  Genus  Quartinia  Ed.  Andre,  1884  (Hymenoptera:  Vespidae: 
Masarinae)  in  Southern  Africa.  Part  I.  Descriptions  of  New  Species  with 

Complete  Venation 

Friedrich  W.  Gess 
Albany  Museum,  Grahamstown,  6140  South  Africa;  email:  f.gess@ru.ac.za 


Abstract. — In  this  publication,  the  first  of  a  projected  series  revising  the  Afrotropical  (essentially 
southern  African)  species  of  the  genus  Quartinia  Ed.  Andre,  1884  (Hymenoptera:  Vespidae, 
Masarinae),  eleven  new  species  are  described.  Of  these,  seven  occurring  variously  in  the  southern 
Namib  Desert  and  in  its  southward  extension  down  the  western  coast  of  South  Africa,  and  one 
occurring  on  the  southern  coast  of  South  Africa,  have  been  found  nesting  in  sand-filled  snail  shells. 
They  are:  australis,  bonaespei,  conchicola,  namaqua,  namaquettsis,  obibensis,  and  refugicoln.  To 
these  species  is  added  vexilhita  which  is  presumed  to  have  the  same  nesting  habits.  A  key  to 
distinguish  these  species  is  given.  The  other  three  newly  described  species,  all  from  Namibia,  are: 
femorata,  geigeriae  and  lamellata. 


Following  van  der  Vecht  and  Carpenter 
(1990)  Quartinia  Ed.  Andre,  1884  is  here 
understood  to  include,  as  junior  subjective 
synonyms,  Quartiniella  Schulthess,  1929 
and  Quartinioides  Richards,  1962. 

As  has  been  pointed  out  by  Carpenter 
(2001),  Quartiniella  and  Quartinioides  were 
primarily  based  on  the  partitioning  of 
a  trend  in  the  reduction  of  wing  venation, 
Quartiniella  being  defined  on  the  basis  of 
the  loss  of  3rs-m  and  2m-cu  and  Quarti- 
nioides because  it  has  2m-eu  present  but 
attenuate  and  interrupted,  whereas  Quarti- 
nia has  it  complete.  In  Quartiniella  in 
particular  and  to  some  extent  also  in 
Quartinioides  reduction  of  wing  venation 
is  a  correlate  of  overall  size  reduction  As 
formal  taxonomic  partitioning  of  essential- 
ly continuous  variation  is  an  unacceptable 
practice,  Carpenter  synonymized  Quarti- 
niella and  Quartinioides  with  Quartinia, 
a  view  with  which  the  present  author  is 
in  full  agreement. 

Nevertheless,  in  view  of  the  large  num- 
ber of  species  in  Quartinia,  adoption  of  the 
above  venational  characters  to  divide  the 
genus  into  smaller,  more  manageable  but 


totally  informal,  non-natural  units  is  found 
to  be  useful.  Thus  the  present  paper  deals 
with  species  with  complete  venation  -  that 
is  species  which  in  the  past  would  have 
been  placed  in  Quartinia  sensu  stricto. 

In  his  revision  Richards  (1962)  dealt  with 
a  total  of  61  southern  African  species,  18 
being  placed  in  Quartinia,  38  in  Quarti- 
nioides and  five  in  Quartiniella.  Of  these,  11, 
26  and  two  respectively  were  described  as 
new.  One  additional  species,  placed  in 
Quartinioides  was  added  (Richards  1982). 
Available  to  Richards  in  1962  were  just 
over  one  thousand  specimens  -  140  Quar- 
tinia, 727  Quartinioides  and  148  Quartiniella. 
Ten  species  were  known  from  only  one 
specimen,  30  species  from  only  one  sex.  It 
is  clear  that  Richards  suffered  from  a  pau- 
city of  material.  Particularly  the  lack  of 
large  samples  from  individual  populations 
spread  over  the  distributional  area  pre- 
vented him  from  appreciating  factors  such 
as  intraspecific  variation  and  geographical 
clines.  In  some  instances  the  associations  of 
sexes  is  of  doubtful  validity,  especially 
where  males  and  females  are  from  widely 
separated  localities. 


212 


Journal  of  Hymenoptera  Research 


The  present  study  is  based  on  over  6000 
specimens,  most  of  which  were  purpose- 
fully collected.  A  large  proportion  of  the 
specimens  have  associated  biological  data 
-  mostly  flower  visiting  records  but  also, 
for  some  species,  nesting  data. 

Desirable  as  it  might  be  to  undertake 
a  complete  revision  of  the  genus,  this  is  at 
present  not  practicable.  Rather  than  to  get 
bogged  down  in  a  study  which  might 
never  be  completed  and  published,  it  is 
intended  to  publish  a  series  of  papers 
describing  new  species  as  well  as  review- 
ing some  known  species.  It  is  envisioned 
that  a  new  key  to  species  will  complete  the 
series. 

Quartinia  species  range  in  length  from 
a  little  over  2  mm  to  7  mm.  In  comparison 
with  the  great  majority  of  species  of  other 
genera  of  Masarinae  even  the  largest 
Quartinia  are  relatively  small.  In  view  of 
the  considerable  range  in  size  shown  by 
species  of  Quartinia  and  in  order  to  express 
relative  size,  categories  based  on  length 
have  been  established  for  species  of  the 
genus.  These  are:  minute  (1.5-2.5  mm); 
small  (2.5-3.5  mm);  medium  (3.5-4.5 
mm);  large  (4.5-5.5  mm);  very  large  (5.5- 
6.5  mm);  and  gigantic  (6.5-7.5  mm). 

The  notation  used  for  expressing  geo- 
graphic co-ordinates  is  as  in  the  gazetteer 
of  The  Times  Atlas  of  the  World  (1981).  The 
figures  before  the  stop  are  degrees,  those 
after  the  stop  are  minutes;  the  stop  is  not 
a  decimal  point. 

For  purposes  of  plotting  distributions, 
co-ordinates  have  been  given  in  square 
brackets  in  the  text  for  those  localities  for 
which  none  are  given  on  the  data  labels. 

On  a  few  data  labels  from  collections 
other  than  that  of  the  Albany  Museum  the 
collecting  locality  is  followed  by  degree 
latitude  and  degree  longitude  and  by  half- 
and  quarter-degree  reference  letters  ac- 
cording to  the  Degree  Reference  System 
of  Leistner  and  Morris  (1976).  As  this 
system  is  not  universally  understood  an 
attempt  has  been  made  here  to  find  on 
a  map  the  localities  concerned  and  to  add 


in  square  brackets  the  co-ordinates  ex- 
pressed in  the  manner  adopted  in  this 
paper. 

In  listing  the  material  examined,  the 
localities  have  been  arranged,  as  far  as 
practicable,  in  north  to  south  order  within 
countries  or,  in  the  case  of  South  Africa, 
within  provinces. 

Acronyms  for  institutions  in  which 
material  is  housed  are:  AMG  =  Albany 
Museum,  Grahamstown,  South  Africa; 
CAS  =  California  Academy  of  Sciences, 
San  Francisco,  United  States  of  America; 
FSCA  =  Florida  State  Collection  of  Arthro- 
pods, Gainesville,  United  States  of  Amer- 
ica; NCP  =  National  Collection  of  Insects, 
Pretoria,  South  Africa;  NNIC  =  Namibian 
National  Insect  Collection,  Windhoek, 
Namibia. 

DESCRIPTION  OF  SPECIES  AND 
COLLECTION  DATA 

A)  Species  nesting  in  sand-filled  snail 
shells  or  (vexillata)  presumed  to  do  so. 

Quartinia  australis  Gess,  new  species 

Diagnosis. — Large  (5.0-5.4  mm).  Fore 
wing  with  Cula  and  2m-cu  complete  and 
as  thick  as  the  other  veins.  Tegula  with 
posterior  inner  corner  inwardly  produced. 
Both  sexes  predominantly  black  with  limit- 
ed white  markings;  male  with  clypeal  disc 
and  underside  of  scape  and  pedicel  white. 

Description. — Female:  Black.  The  follow- 
ing are  white:  narrow  anterior  margin  of 
pronotum  (in  most  specimens)  and  extreme 
postero-dorsal  angle  of  same;  tegula  anteri- 
orly and  posteriorly;  lateral  lamella  of 
scutellum;  posterior  bands  medially  on 
terga  I-V  (that  on  V  in  some  specimens 
reduced  to  a  postero-medial  spot);  distal 
end  of  fore  femur;  streaks  on  fore  and 
middle  tibiae;  proximal  and  distal  ends  of 
hind  tibia.  Brown  are:  rest  of  legs;  underside 
of  flagellum.  Wings  lightly  browned. 

Length  5.0-5.4  (average  of  5:5.3  mm); 
length  of  fore  wing  3.4-3.6  mm  (average 
of  4:3.53  mm);  hamuli  6. 


Volume  16,  Number  2,  2007 


213 


Head  in  front  view  1.31  X  as  wide  as 
long,  finely  microreticulate,  matt;  clypeus 
apunctate;  frons  and  vertex  with  shallow 
punctures  separated  by  about  their  width 
(punctures  barely  perceivable  on  lower 
regions  of  frons,  clearer  in  ocular  sinuses 
and  upper  part  of  frons  and  particularly  on 
vertex.  POL:OOL  =  1:0.6.  Clypeus  1.5  X  as 
wide  as  long;  anterior  margin  shallowly 
and  widely  emarginate;  antero-lateral  an- 
gles rounded. 

Mesosoma  microreticulate,  moderately 
shiny,  with  punctures  larger  and  more 
obvious  than  on  head. 

Gaster  microreticulate  but  shiny;  punc- 
tures finer  and  shallower  than  on  head  and 
much  more  so  than  those  on  mesosoma, 
becoming  progressively  finer  posteriorly. 

Male:  Black.  White  markings  as  in  fe- 
male, with  in  addition:  labrum;  disc  of 
clypeus  (i.e.  not  sides  nor  areas  adjacent  to 
antennal  sockets);  underside  of  scape  and 
pedicel;  posterior  band  on  tergum  VI; 
distal  end  of  middle  and  hind  femora. 
Underside  of  flagellum  white  suffused 
with  reddish-brown. 

Length  5.0-5.4  (average  of  4:5.1  mm); 
length  of  front  wing  3.4-3.6  mm  (average 
of  4:3.42  mm);  hamuli  6. 

Head  in  front  view  1.5  X  as  wide  as  long; 
POL:OOL  =  1:0.6.  Clypeus  1.5  X  as  wide 
as  long;  anterior  margin  shallowly  and 
widely  emarginate;  antero-lateral  angles 
rounded. 

Microsculpture  and  punctuatation  of 
head  and  body  similar  to  that  of  female. 

Tergum  VII  (Fig.  6)  with  surface  notice- 
ably depressed  and  with  hindmargin  with 
a  short  median  slit.  Sterna  I-VI  unmodi- 
fied; sternum  VII  trilobed,  basally  marked- 
ly concave  between  swollen  and  poster- 
iorly produced  lateral  lobes  and  with 
median  lobe  flat  and  densely  setose. 

Etymology. — The  name  australis  draws 
attention  to  the  southern  provenance  of 
the  species. 

Material  examined.— Holotype:  J,  SOUTH 
AFRICA:   WESTERN   CAPE:   Witsand    (34.23S 


20.52E),  14.viii.2002  (F.  W.  and  S.  K.  Gess)  (ex 
nest  in  shell  of  Tlieba  pisana  (Mull.),  Helicidae) 
[AMG].  Paratypes:  SOUTH  AFRICA:  WEST- 
ERN CAPE:  same  data  as  holotype,  7  99,  4  Jg 
(ex  nests  in  shells  of  Tlieba  pisana  (Mull.), 
Helicidae)  [AMG]. 

Geographic  distribution. — Known  only 
from  the  type  locality,  Witsand,  near  Port 
Beaufort  at  the  mouth  of  the  Bree  River, 
a  little  to  the  west  of  the  southernmost 
point  of  Africa. 

Floral  associations. — Unknown. 

Nesting.— Found  nesting  in  sand-filled 
shells  of  the  exotic  Tlieba  pisana  (Mull.) 
(Mollusca:  Gasteropoda:  Pulmonata:  Heli- 
cidae) collected  from  the  surface  of  the 
sand  below  bushes  growing  on  supra- 
littoral  dunes. 

Quartinia  bonaespei  Gess,  new  species 

Diagnosis.— Very  large  to  gigantic  (6.3- 
7.0  mm).  Fore  wing  with  Cula  and  2m~cu 
complete  and  as  thick  as  other  veins. 
Tegula  short,  laterally  rounded,  with  pos- 
terior inner  corner  inwardly  produced. 
Both  sexes  black  with  white-marked  pro- 
notum,  tegula  and  scutellar  lamella  and 
with  wide,  bright  reddish-orange  posterior 
bands  on  all  but  last  two  terga.  Male  with 
greatly  enlarged  and  modified  fore  leg, 
with  somewhat  modified  middle  and  hind 
legs,  with  tergum  VII  apico-medially 
drawn  out  into  a  robust,  dorsally  flattened 
and  apically  rounded  process,  and  with 
sterna  medially  depressed. 

Description. — Female:  Black.  The  follow- 
ing are  white:  medially  interrupted  trans- 
verse band  on  dorsum  of  pronotum  and 
minute  dot  at  postero-dorsal  angle  of  same; 
anterior  and  posterior  thirds  of  tegula 
(median  third  black);  medially  broadly 
interrupted  band  on  lamellate  margin  of 
scutellum.  Bright  reddish-orange  are: 
mandibles  distally;  posterior  markings 
dorsally  (i.  e.  not  extending  down  sides) 
on  terga  I-IV  (that  of  tergum  I  wide, 
covering  entire  dorsal  surface,  that  of  II 
slightly  narrower,  that  of  III  wide  medially 
but  narrowed  laterally,  that  oi  IV  a  median 


214 


Journal  of  Hymenoptera  Research 


transverse  spot).  Labrum  brown.  Under- 
side of  antennae,  to  various  degrees,  pale. 
Coxa,  trochanter,  femur  and  tibia  of  all  legs 
black  with  exception  of  yellowish  streak  on 
antero-dorsal  surface  of  fore  tarsus  and 
same  colour  on  extreme  base  of  middle  and 
hind  tibiae;  tarsomeres  dark  brown.  Wings 
brown;  veins  dark  brown  to  black. 

Length  6.3-7.0  mm  (average  of  5:6.7 
mm);  length  of  fore  wing  4.3-4.5  mm 
(average  of  5:4.4  mm);  hamuli  6. 

Head  in  front  view  1.29  X  as  wide  as 
long,  microreticulate,  matt,  with  small, 
shallow  punctures  (sparse  on  clypeus,  well 
separated  on  lower  part  of  frons  but 
progressively  closer  on  upper  part  of  frons 
and  on  vertex).  POL:OOL  =  1:0.75.  Clyp- 
eus 1.3  X  as  wide  as  long;  anterior  margin 
shallowly  emarginate;  antero-lateral  angles 
rounded. 

Mesosoma  microreticulate,  matt,  with 
punctures  slightly  larger  and  deeper  than 
on  head  (moderately  well  separated  on 
mesoscutum  and  scutellum,  closer  on  pro- 
notum  and  upper  part  of  mesopleuron 
where  sculpture  almost  reticulate-punctate). 
Gaster  microreticulate  but  shiny;  punc- 
tures finer  and  shallower  than  on  head  and 
mesosoma,  becoming  progressively  finer 
posteriorly. 

Male:  Black.  White  markings  as  in  fe- 
male. Bright  reddish-orange  markings  on 
gaster  similar  to  those  of  female  but 
present  also  on  tergum  V  where  transverse 
as  on  anterior  terga.  Underside  of  flagello- 
meres,  antero-distal  spot  on  fore  femur, 
dorsal  and  anterior  surfaces  of  fore  tibia, 
fore  tarsus,  yellowish-orange. 

Length  6.3  mm;  length  of  fore  wing 
4.6  mm;  hamuli  6. 

Head  in  front  view  1.33  X  as  wide  as 
long,  much  more  finely  microreticulate  and 
much  more  finely  punctate  than  in  female, 
moderately  shiny.  POL:OOL  -  1:0.7.  Clyp- 
eus shorter  than  that  of  female,  1.46  X  as 
wide  as  long. 

Mesosoma  much  more  finely  microreti- 
culate and  much  more  finely  punctate  than 
in  female,  moderately  shiny. 


Fore  leg  much  modified;  coxa  and 
trochanter  enlarged;  femur  (Fig.  1)  greatly 
swollen,  postero-basally  with  pointed  tu- 
bercle, its  posterior  surface  depressed, 
smooth  and  very  shiny  and  forming  an 
angle  with  ventral  surface;  tibia  greatly 
enlarged,  ventrally  with  its  swollen  basal 
section  fitting  into  opposing  disto-ventral 
emargination  of  femur  (best  seen  in  ante- 
rior view);  tarsomeres  robust,  noticeably 
setose.  Middle  and  hind  legs  more  robust 
than  those  of  female;  femora  of  both  these 
legs  swollen  beneath  but  longitudinally 
grooved  over  distal  half  to  accommodate 
tibia  when  opposed;  tarsomeres  II— IV  of 
middle  leg  noticeably  wider  than  those  of 
hind  leg. 

Gaster  very  finely  microreticulate,  shiny. 

Tergum  VII  (Fig.  7)  baso-laterally  with 
a  pronounced  rounded  tubercle,  apico- 
medially  drawn  out  into  a  robust,  dorsally 
flattened  and  apically  rounded  process 
raised  above  depressed  surface  on  either 
side  of  it;  process  dorsally  with  a  slight 
median  longitudinal  carina  and  laterally  on 
each  side  with  a  smooth  low  carina  (carried 
forward  some  distance  onto  the  tergal  disk) 
at  angle  formed  by  its  dorsal  and  lateral 
surfaces;  hind  margin  of  tergum  in  lateral 
view  forming  a  low  smooth  curve  from 
basal  tubercle  to  tip  of  apical  process. 

Sternum  II— VI  depressed  medially;  ster- 
num II  markedly  so;  III— VI  progressively 
less  so. 

Etymology. — The  name  bonaespei,  a  Latin 
noun  in  the  genitive,  refers  to  the  Cape  of 
Good  Hope  and  draws  attention  to  the 
provenance  of  the  species,  especially  to  the 
type  locality  which  is  within  sight  of  Table 
Mountain. 

Material  examined. — Holotype:  o,  SOUTH 
AFRICA:  WESTERN  CAPE:  on  coast  4  km 
north  of  Bloubergstrand  (33.46S  18.27E),  12- 
13.viii.2002  (F.  W.  and  S.  K.  Gess)  (on  ground) 
[AMG].  Paratypes:  SOUTH  AFRICA:  WEST- 
ERN CAPE:  Donkinsbaai,  10  km  S  of  Door- 
nbaai,  low  vegetated  dunes  and  slacks  behind 
beach  (31.54S  18.17E),  9.ix.2005  (F.  W.  and  S.  K. 
Gess),  8  99  (4  99  from  sand-filled  Trigonephrus 


Volume  16,  Number  2,  2007 


215 


Figs.  1-5.     Left  fore  femur  of  male  (posterior  view).  1.  Quartinia  bonaespei,  2.  Quartinia  conchicola,  3.  Quartinia 

namaquensis,  4.  Quartinia  vexillata,  5.  Quartinia  fenwrata. 


shells;  2  99  reared  from  mature  larvae  ex 
Quartinia  nests  ex  sand-filled  Trigouephrus 
shells,  emerged  in  lab.  first  week  of  June  2006; 

2  99  visiting  white  centred,  pink  flowers  of 
Drosantheinuin  sp.,  Aizoaceae:  Mesembryan- 
thema)  [AMG];  Lamberts  Bay,  dunes  behind 
beach  (32.05S  18.19E),  28.ix.2005  (F.  W.  and  S.  K. 
Gess),  1  9  (from  Trigonepihrus  shell)  [AMG]; 
Lamberts  Bay,  sandy  southern  bank  of  lagoon 
(32.05S  18  19E),  28.ix.2005  (F.  W.  and  S.  K.  Gess), 

3  99  (visiting  yellow  flowers  of  Conicosia, 
Aizoaceae:  Mesembryanthema)  [AMG];  Elands 
Bay,  low  vegetated  dunes  behind  beach  (32.19S 
18.20E),  30.ix.2005  (F.  W.  and  S.  K.  Gess),  3  99,  1 
J  (1  9  from  sand  filled  Trigouephrus  shell;  2  99 
visiting  pink  flowers  of  Drosanthemum,  Aizoa- 
ceae: Mesembryanthema;  1  ,_J  reared  ex  Quarti- 
nia nest  in  sand-filled  Theba  pisana  (Mull.)  shell, 
emerged  in  lab.  6.viii.2006)  [AMG];  Roscherpan 
Nature  Reserve  (32.36S  18.18E),  24.iii.2001 
(Feuerer  &  Thell),  4  99,   1    J   (from  shells  of 


Trigouephrus  porphyrostoma  (Melvill  &  Pon- 
sonby)  [Zool.  Mus  Berlin];  Yzerfontein  (33.20S 
18.10E),  15.X.2006  (D.  W.,  G.  T.  and  G.  M.  Gess), 
1  9  (ex  Theba  pisana  shell)  [AMG];  S  of 
Yzerfontein  (33.22S  18.1  IE),  15.X.2006  (D.  W., 
G.  T.  and  G.  M.  Gess),  1  9  (on  sand)  [AMG]; 
Melkbosstrand  (33.42S  18.26E),  lO.x.2005  (F.W. 
and  S.  K.  Gess),  2  99  (1  9  on  sand  beneath 
flowering  Tracln/andra  divaricata  (Jacq.)  Kunth., 
Asphodelaceae;  1  9  reared  from  mature  larva  ex 
Quartinia  nest  ex  sand-filled  Theba  pisana  shell) 
[AMG];  on  coast  4  km  north  of  Bloubergstrand 
(33.46S  18.27E),  12-13.viii.2002  (F.  W.  and  S.  K. 
Gess),  11  99,  (6  99,  3  99  visiting  white  flowers  of 
Tracln/andra  divaricata;  1  9  visiting  purplish  pink 
flowers  of  Aizoaceae:  Mesembryanthema;  1  9  ex 
nest  in  sand-filled  Trigouephrus  shell)  [AMG]; 
same  locality,  5.X.2005  (F.  W.  and  S.  K.  Gess),  4 
99, 1  6*  (1  9  from  sand  filled  Trigouephrus  shell;  1 
9  visiting  white  flowers  of  Tracln/andra  divar- 
icata; 2  99  on  sand  beneath  flowering  Tracln/au- 


216 


Journal  of  Hymenoptera  Research 


6. 


8. 


10. 


11 


12. 

Figs.  6-13.  Tergum  VII  of  male  (postero-dorsal  view).  6.  Quartinia  australis,  7.  Quartinia  bonaespei,  8.  Qumiinia 
conchkola,  9.  Quartinia  namaquensis,  10.  Quartinia  obibensis,  11.  Quartinia  refugicola,  12.  Quartinia  vcxillata,  13. 
Quartinia  femorata.  [TergumVII  of  Quartinia  namaqua  is  very  similar  to  that  of  Quartinia  obibensis  (Fig.  10)]. 


dm  divaricata;  1  j  reared  ex  Quartinia  nest  in 
sand-filled  Theba  pisana  shell,  emerged  in  lab. 
8.viii.2006)  [AMG]. 

Geographic  distribution. — Known  only 
from  the  supra-littoral  dunes  of  the  Atlan- 
tic seaboard  of  the  Western  Cape,  from 
Donkinsbaai,  circa  220  km  north  of  Cape 
Town  to  Bloubergstrand  at  the  northern 
extremity  of  Table  Bay  (the  type  locality). 
At  Yzerfontein  it  has  been  found  together 
with  Q.  namaqua  and  Q.  obibensis. 

Floral  associations. — Asphodelaceae  (Tra- 
chyandra)  and  Aizoaceae:  Mesem- 
bryanthema  (including  Conicosia  and  Dro- 
santhemum). 


Nesting. — The  collection  at  all  the  listed 
localities  of  adult  females  from  sand-filled 
snail  shells,  the  discovery  of  an  adult 
female  at  Bloubergstrand  in  a  shell  contain- 
ing also  an  open  cell  provisioned  with 
a  mixture  of  pollens  including  that  of 
Trachyandra  divaricata,  and  the  rearing  in 
the  lab  of  adults  from  mature  larvae 
extracted  from  cells  found  in  shells  from 
four  of  the  localities,  demonstrates  that  this 
species,  like  others  occurring  in  sandy 
areas,  utilizes  sand-filled  snail  shells  as 
a  nesting  niche.  Shells  of  the  indigenous 
desert  snail,  Trigonephrus  species  (Mol- 
lusca:  Gasteropoda:   Pulmonata:   Dorcasii- 


Volume  16,  Number  2,  2007 


217 


dae)  are  the  original  ones  utilized  and 
appear  to  be  preferred;  where  these  are  in 
short  supply,  the  smaller,  thinner  and 
therefore  less  opaque  shells  of  the  exotic 
Theba  pisana  (Mull.)  (Mollusca:  Gastero- 
poda: Pulmonata:  Helicidae)  are  used. 

Quartinia  conchicola  Gess,  new  species 

Quartinia  sp.  (larger  sp.)  (Gess  and  Gess  1999, 
nesting) 

Diagnosis. — Very  large  (5.6-6.3  mm). 
Fore  wing  with  Cula  and  2m-cu  complete 
and  as  thick  as  other  veins.  Both  sexes  with 
vertex  behind  posterior  ocelli  depressed, 
somewhat  concave;  with  fore  coxa  not 
swollen  basally  nor  anteriorly  produced 
but  evenly  curved.  Male  with  fore  femur 
enlarged,  excavated  beneath  and  undulate 
postero-ventrally;  tibia  robust,  markedly 
swollen,  appreciably  shorter  than  femur 
and,  when  opposed  to  femur,  fitting  into 
ventral  excavation  of  same. 

Description. — Female:  Black.  The  follow- 
ing are  yellowish-white:  short  (almost 
medially  interrupted)  and  laterally  widen- 
ing transverse  band  on  dorsum  of  prono- 
tum  and  minute  spot  at  postero-dorsal 
angle  of  same;  humeral  streak  of  varying 
length;  anterior  and  posterior  thirds  of 
tegula  (median  third  clear,  testaceous); 
medially  interrupted  band  on  lamellate 
margin  of  scutellum  (specimens  from 
Hondeklip  Bay  only).  (The  specimen  from 
Knersvlakte  lacks  the  humeral  streak  as  do 
those  from  between  Alexander  Bay  and 
Port  Nolloth  which  in  addition  have  the 
other  markings  on  the  thorax  reduced  and 
reddish-brown.  Those  from  W  of  Wallek- 
raal  are  without  thoracic  markings.)  The 
following  are  various  shades  of  light 
reddish  brown:  mandible  (other  than  base); 
labrum;  lower  aspect  of  pedicel  and  flagel- 
lum;  posterior  bands  (in  some  specimens 
widened  medially  and  usually  not  attain- 
ing lateral  margins)  on  terga  I-IV  (or  V); 
apices  of  all  femora;  most  of  tibia  and 
tarsus  of  all  legs.  Venation  light  brown  at 
base  of  wings,  otherwise  very  dark  brown. 


Wing  membrane  very  slightly  browned. 
Length  5.6-6.3  mm  (average  of  8  = 
6.1  mm);  length  of  fore  wing  3.7-4.2  mm 
(average  of  8  =  4.1  mm);  hamuli  6. 

Head  in  front  view  1.3-1.34  X  as  wide  as 
long;  POL:OOL  =  1:0.65  (average  of  5). 
Vertex  behind  posterior  ocelli  depressed, 
somewhat  concave. 

In  general  facies  similar  to  male  (de- 
scribed below)  but  with  legs  and  last 
tergum  simple. 

Male:  Head  and  mesosoma  black,  gaster 
and  greater  part  of  femora  of  all  legs  very 
dark  brown  to  almost  black.  The  following 
are  yellowish-white:  pair  of  small  spots  on 
frons  immediately  above  frontoclypeal  su- 
ture (in  specimens  from  north  of  Vanrhyns- 
dorp  only);  short  (almost  medially  inter- 
rupted) and  laterally  widening  transverse 
band  on  dorsum  of  pronotum  and  minute 
spot  at  postero-dorsal  angle  of  same;  hu- 
meral streak  of  varying  length;  anterior  and 
posterior  thirds  of  tegula  (median  third 
clear,  testaceous);  medially  interrupted 
band  on  lamellate  margin  of  scutellum.  (In 
a  specimen  from  between  Alexander  Bay 
and  Port  Nolloth  the  humeral  markings  are 
absent  and  the  other  markings  on  the  thorax 
are  reddish-brown.)  The  following  are 
various  shades  of  light  reddish  brown: 
mandible  (other  than  base);  labrum;  lower 
aspect  of  scape,  pedicel  and  flagellum; 
posterior  bands  (slightly  widened  medially 
and  laterally  but  not  attaining  lateral  mar- 
gins) on  terga  I— VI;  apices  of  all  femora; 
most  of  fore  tibia;  middle  tibia  and  hind 
tibia  to  variable  extent  and  tarsus  of  all  legs. 
Venation  light  brown  at  base  of  wings, 
otherwise  very  dark  brown.  Wing  mem- 
brane very  slightly  browned. 

Length  circa  5.8-6  mm;  length  of  fore 
wing  circa  4^4.5  mm. 

Head,  mesosoma  and  terga  I— VII  very 
finely  microsculptured  (shagreened)  but 
nevertheless  shiny. 

Head  in  front  view  1.4-1.45  X  as  wide  as 
long;  POL:OOL  =  1:0.65.  ).  Vertex  behind 
posterior  ocelli  depressed,  somewhat  con- 
cave. 


218 


Journal  of  Hymenoptera  Research 


Tegula  with  posterior  inner  corner  in- 
wardly produced.  Wing  venation  with  Cul 
and  2m-cu  complete  and  as  thick  as  other 
veins. 

Fore  leg  with  coxa  unmodified;  femur 
(Fig.  2)  enlarged,  excavated  beneath  and 
undulate  postero-ventrally;  tibia  robust, 
markedly  swollen,  appreciably  shorter 
than  femur  and  when  opposed  to  femur 
fitting  into  ventral  excavation  of  same. 

Middle  and  hind  femora  robust  but 
otherwise  not  markedly  modified. 

Sternum  I  postero-medially  very  slightly 
bi-tuberculate;  sternum  II  somewhat  raised 
on  either  side  of  median  area.  Tergum  VII 
(Fig.  8)  in  posterior  half  with  dorsal  surface 
raised  laterally  and  delimited  by  low 
carinae,  produced  apically  and  with  a  deep, 
narrow,  slightly  sub-parallel  median  slit. 

Etymology. — The  name  conchicola  is 
a  compound  word  formed  from  the  Latin 
words  concha  -  ae  -  the  shell  of  a  mollusc, 
and  cola  -  a  dweller.  It  serves  to  draw 
attention  to  the  species'  association,  albeit 
not  unique,  with  sand-filled  shells  of  the 
Desert  Snail,  Trigonephrus,  in  which  its 
nests  are  sheltered  from  prevailing  winds. 

Material  examined.— Holotype:  J,  SOUTH 
AFRICA:  WESTERN  CAPE:  12  km  N  of  Vanr- 
hynsdorp  (31.31S  18.43E),  27.ix.2005  (F.  W.  and 
S.  K.  Gess)  (dead,  ex  nest  in  sand-filled 
Trigonephrus  shell)  [AMG].  Paratypes:  SOUTH 
AFRICA:  NORTHERN  CAPE:  Richtersveld  Na- 
tional Park,  1.5  km  from  Helskloof  Gate  (28.18S 
16.57E),  8.ix.l996  (F.  W.,  S.  K.  and  R.  W.  Gess),  1 
9  (on  white  flowers  of  Pelargonium  klinghardtense 
Knuth,  Geraniaceae)  [AMG];  Richtersveld,  W  of 
Brandkaros  (28.29S  16.40E),  15.ix.1996  (F.  W.,  S. 
K.  and  R.  W.  Gess),  1  S  (dead)  and  fragments  of 
2  further  <$<$  (ex  nests  in  sand-filled  Trigonephrus 
shells)  [AMG];  between  Alexander  Bay  and 
turnoff  to  Oranjemund  (28.35S  16.30E) 
13.ix.1996  (F.  W.,  S.  K.  and  R.  W.  Gess),  1  9 
(dead,  ex  nest  in  sand-filled  Trigonephrus  shell) 
[AMG];  24  km  S  of  Alexander  Bay  on  road  to 
Port  Nolloth  [=  60  km  N  of  Port  Nolloth  on 
road  to  Alexander  Bay]  (28.47S  16.38E), 
27.ix.1997  (F.  W.  and  S.  K.  Gess),  5  99  (4  ex 
nests  in  sand-filled  Trigonephrus  shells;  1  on 
ground)  [AMG];  same  locality,  ll.x.2000  (F.  W. 


and  S.  K.  Gess),  1  9  (visiting  pink  flowers  of 
Drosanthemum  sp.)  [AMG];  60  km  S  of  Alexan- 
der Bay  on  road  to  Port  Nolloth  (28.51  S  16.40E), 
19.ix.1996  (F.  W.,  S.  K.  and  R.  W.  Gess),  1  J 
(dead,  ex  nest  in  sand-filled  Trigonephrus  shell) 
[AMG];  Hondeklip  Bay  (30.19S  17.17E), 
12.X.1994  (F.  W.  and  S.  K.  Gess),  3  99  (visiting 
yellow  flowers  of  Conicosia  sp.,  Aizoaceae: 
Mesembryanthema)  [AMG];  W  of  Wallekraal 
(30.21S  17.26E),  8.X.1997  (F.  W.  and  S.  K.  Gess),  2 
99  (live)  and  fragments  of  1  c?  (ex  nests  in  sand- 
filled  Trigonephrus  shells)  [AMG].  WESTERN 
CAPE:  Knersvlakte,  48  km  N  of  Vanrhynsdorp 
(31.14S  18.32E),  20.ix.1996  (F.  W.,  S.  K.  and  R.  W. 
Gess),  1  9  [AMG];  12  km  N  of  Vanrhynsdorp 
(31.31S  18.43E),  27.ix.2005  (F.  W.  and  S.  K.  Gess), 
2  99,  2  S3  (dead  and  incomplete,  ex  nests  in 
sand-filled  Trigonephrus  shells)  [AMG];  SE  of 
Lutzville  on  road  to  Vredendal  (31.36S  18.23E), 
29.ix.2005  (F.  W.  and  S.  K.  Gess),  399  (reared 
from  mature  larvae  ex  Quartinia  nests  ex  sand- 
filled  Trigonephrus  shells,  emerged  in  lab  at  the 
end  of  April  2007)  [AMG]. 

Geographic  distribution. — The  species  is 
known  from  South  Africa  from  the  western 
part  of  the  Northern  Cape,  mainly  along 
the  seaboard  from  the  Orange  River  south- 
wards, and  from  the  northwestern  Western 
Cape  where  it  extends  inland  to  a  distance 
of  about  50  km.  In  occurs  variously  togeth- 
er with  Q.  namaqua,  Q.  namaquensis,  Q. 
obibensis,  Q.  rufigicola  and  Q.  vexillata. 

Floral  associations. — Aizoaceae:  Mesem- 
bryanthema (Conicosia,  Drosanthemum), 
Geraniaceae  (Pelargonium). 

Nesting. — Throughout  its  presently 
known  distributional  area  found  nesting 
in  sand-filled  shells  of  the  desert  snail 
Trigonephrus  sp.  (Mollusca:  Gasteropoda: 
Pulmonata:  Dorcasiidae).  For  further  de- 
tails see  Gess  and  Gess  (1999). 

Discussion. — Q.  conchicola  and  Q.  vexillata 
appear  to  be  closely  allied  and  at  least  in 
the  north-western  Richtersveld  (S  of  Alex- 
ander Bay)  overlap  in  their  distribution. 
Whereas  the  males  are  readily  distinguish- 
able on  the  basis  of  secondary  sexual 
characters  -  notably  the  differently  modi- 
fied fore  legs  -  the  females  are  deceptively 
similar  and  at  first  sight  are  very  difficult 
to  separate.  They  may,  however,  be  distin- 


Volume  16,  Number  2,  2007  219 

guished  by  characters  which   they  share  (progressively  darkened)  of  all  legs;  in  two 

with  their  respective  males:  Q.  conchicola  by  of  the  southern  specimens  apex  of  femur, 

the  depressed,  somewhat  concave  vertex  base  and  apex  of  tibia  and  base  of  first 

and  by  the  unmodified  fore  coxae  and  Q.  tarsomere  only.  Wings  slightly  darkened; 

vexillata  by  the  evenly  convex  vertex  and  veins  brown. 

by  the  basally  swollen  and  anteriorly  pro-  Length   6.2  mm;    length    of   fore    wing 

duced  fore  coxae.  3.9  mm;  hamuli  6. 

Head  in  front  view  1.3  X  as  wide  as  long; 

Quartinia  namaqua  Gess,  new  species  clypeus   1.1    x    as   wide  as   long   (length 

measured    to    bottom    of   emargination); 

Diagnosis.-Very    large    (5.8-6.2  mm).  pOL:OOL         1:09    Q                       dosel 

Fore  wing  with  Cula  and  2m-cu  complete  microsculptured/  with  bareiy  discernable 

and  as  thick  as  other  veins.  Tegula  with  shaUow           tureS/  dull;  frons  and  vertex 

posterior  inner  corner  markedly  inwardly  similarly  micr0sculptured  but  somewhat 

produced,  reddish  brown.  Both  sexes  with  mQre   obviously   punctured,    moderately 

head  and   thorax  black  (except,  in  most  shiny;    mesosoma    micr0sculptured    with 

specimens,  a  small  reddish-brown  marking  obvious  shallow  pictures;  interstices  of 

medially  on  anterior  margin  of  pronotum);  puncture  width  or  less;  parapsidal  furrows 

gaster  black  with  a  variable  number  of  very   obvious;   gaster   finely   and   closely 

reddish  brown  posterior  bands  which  do  punctured  shinv 

not  attain  sides  of  terga.  Parapsidal  fur-  Mak.  Black  The  foUowing  are  reddish- 
rows  very  noticeable.  Male  with  clypeus  brown:  mandibles  distally;  scape  apically, 
evenly  convex  (not  medially  depressed),  pedical/  upper  and  iower  side  0f  flagello- 
closely  and  finely  sculptured;  with  ster-  meres  (except  dista]  part  0f  dub);  trans- 
num  I  postero-medially  raised  into  a  pro-  verse  marking  on  anterior  margin  of 
nounced  tubercle;  tubercle  viewed  from  pronotum;  tegula;  posterior  band  not  at- 
behind  with  widely  rounded  (almost  sub-  taining  sides  on  tergum  I  and  mere  in- 
truncate)  apex,  viewed  from  the  side  dication  of  band  on  tergum  II;  apex  of 
sloping  steeply  anteriorly  and  falling  steep-  femur/  most  of  tibia,  tarsomeres  (progres- 
ly  posteriorly  to  hind  margin  of  sternum.  siveiy  darkened)  of  all  legs. 
Tergum  VII  with  distinct  dorsal  and  lateral  Length  5.8  mm;  length  of  fore  wing 
surfaces;  apex  drawn  out  into  a  pair  of  35  mm;  hamuli  6. 

parallel  processes  flanking  narrow  and  slit-  Head  in  front  view  1.4  x  as  wide  as  long; 

like  emargination;  emargination  produced  clypeUs   1.1    X    as   wide   as   long   (length 

anteriorly  as  a  median  impression.  measured    to    bottom    of   emargination); 

Description.— Female:  Black.  The  follow-  POL:OOL  =  1:0.8.  Clypeus  evenly  convex, 

ing  are  reddish-brown:  mandibles  distally;  closely  microsculptured,  with  barely  dis- 

underside  of  pedicel  and  flagellum;  in  two  cernable  shallow  punctures,  only  moder- 

of  the  northern  specimens  a  mere  indica-  ately   shiny;    frons   and    vertex   similarly 

tion  of  a  transverse  marking  on  anterior  microsculptured  but  somewhat  more  obvi- 

margin    of   pronotum;    tegula;    posterior  ously  punctured,  moderately  shiny;  meso- 

bands  not  attaining  sides  on  terga   1-IV  soma  microsculptured  with  obvious  shal- 

(in  southern  specimens  on  terga  I — III  only);  low    punctures;    interstices    of   puncture 

that  on  I  of  even  width  and  covering  about  width   or   less;   parapsidal    furrows   very 

half  of  tergum;  those  of  terga  II  and   III  obvious;  gaster  finely  and   closely  punc- 

narrower  but  medially  expanded;  that  of  tured,  shiny.  Sternum  I  postero-medially 

IV   short   or   (in    two   specimens)    barely  raised  into  a  pronounced  tubercle;  tubercle 

indicated;  in  northern  specimens  apex  of  viewed  from  behind  with  widely  rounded 

femur,  entire  or  most  of  tibia,  tarsomeres  (almost  subtruncate)   apex,   viewed    from 


220 


Journal  of  Hymenoptera  Research 


the  side  sloping  steeply  anteriorly  and 
falling  steeply  posteriorly  to  hind  margin 
of  sternum.  Tergum  VII  with  distinct 
dorsal  and  lateral  surfaces;  apex  drawn 
out  into  a  pair  of  parallel  processes  flank- 
ing narrow  and  slit-like  emargination; 
emargination  produced  anteriorly  as  a  me- 
dian impression. 

Etymology.— The  name,  namaqua,  a  noun 
in  apposition  to  the  generic  name,  is  derived 
from  the  Namaqua  people  of  Namaqualand 
and  refers  to  the  provenance  of  the  species. 

Material  examined.  ■  Holotype,  6\  SOUTH 
AFRICA:  NORTHERN  CAPE:  Inland  of  Hon- 
deklip  Bay  (30.19S  17.17E),  25.ix.2005  (F  W  and  S 
K  Gess)  (ex  nest  in  sand-filled  Trigonephrus 
shell)  [AMG].  Paratypes:  SOUTH  AFRICA: 
NORTHERN  CAPE:  same  data  as  holotype,  1 
9  [AMG];  between  Hondeklip  Bay  and  Wallek- 
raal  (30.22S  17.28E),  25.ix.2005  (F  W  and  S  K 
Gess),  1  9  (ex  nest  in  sand-filled  Trigonephrus 
shell)  [AMG].  WESTERN  CAPE:  SE  of  Lutzville 
on  road  to  Vredendal  (31.36S  18.23E),  29.ix.2005 
(F  W  Gess  and  S  K  Gess),  1  9  (visiting  yellow 
flowers  of  Conicosia  spv  Aizoaceae:  Mesem- 
bryanthema)  [AMG];  Yzerfontein  (33.20S 
18.10E),  15.X.2006  (D.  W.,  G.  T.  and  G.  M.  Gess), 
3  99  (ex  Theba  pisana  shells)  [AMG]. 

Geographic  distribution. — The  species  is 
known  from  South  Africa  from  the  south- 
western Northern  Cape,  from  the  north- 
western Western  Cape,  and  from  Yzerfon- 
tein in  the  southwestern  Western  Cape, 
and  therefore  will  probably  be  found  to 
occur  all  along  the  coastal  sandveld  be- 
tween the  above  areas.  In  occurs  variously 
together  with  Q.  bonaespei,  conchicola,  na- 
maquensis  and  obibensis. 

Floral  associations. — Aizoaceae:  Mesem- 
bryanthema  (Conicosia). 

Nesting. — At  two  localities  found  nesting 
in  sand-filled  shells  of  the  desert  snail 
Trigonephrus  sp.  (Mollusca:  Gasteropoda: 
Pulmonata:  Dorcasiidae)  and  at  another 
obtained  from  shells  of  the  exotic  Tlieba 
pisana  (Mull.)  (Mollusca:  Gasteropoda: 
Pulmonata:  Helicidae). 

Discussion. — Q.  namaqua  is  superficially 
very  similar  to  Q.  obibensis,  most  notably  in 


the  male  in  the  possession  of  a  raised 
tubercle  postero-medially  on  sternum  I.  It 
may  be  distinguished  in  both  sexes  by  the 
more  distinct  and  somewhat  less  close 
puncturation  of  the  mesoscutum  and  scu- 
tellum,  by  the  broader  and  much  more 
noticeable  parapsidal  furrows,  and  by  the 
interocellar  distance  only  slightly  exceed- 
ing the  ocellar-ocular  distance  [POL:OOL 
=  1:0.9  (9)  and  1:0.8  (S)  as  against  1:0.7 
(both  sexes)].  The  male  may  be  distin- 
guished by  the  evenly  convex,  closely 
microsculptured  and  only  moderately 
shiny  clypeus  in  contrast  to  the  medially 
depressed,  non-microsculptured  but 
sparsely  punctured  and  shiny  clypeus  of 
Q.  obibensis.  In  colour  pattern  the  species 
differs  in  that  the  reddish-brown  markings 
are  reduced,  most  notably  in  that  the 
posterior  bands  on  the  gaster  do  not  attain 
the  lateral  margins  of  the  terga. 

Quartinia  namaquensis  Gess,  new  species 

Diagnosis. — Very  large  (5.8-6.0  mm). 
Fore  wing  with  Cula  and  2m-cu  complete 
and  as  thick  as  other  veins.  Tegula  short, 
laterally  rounded,  with  posterior  inner 
corner  inwardly  produced.  Male  black 
with  white-marked  labrum,  clypeus,  frons, 
pronotum,  tegula,  scutellar  lamella,  and 
terga  I-VI.  Fore  leg  greatly  enlarged  and 
modified;  middle  and  hind  legs  somewhat 
modified.  Tergum  VII  drawn  out  apico- 
medially  into  a  robust,  pointed,  dorsally 
flattened  and  apically  narrowly  rounded 
process. 

Description. — Male:  Black.  The  following 
are  white:  labrum;  disc  of  clypeus;  parao- 
cular  streak  from  mandibular  insertion  to 
level  of  top  of  antennal  socket  (specimen 
from  Wallekraal  only);  supra-clypeal  mark- 
ing (more  or  less  quadrate  and  bilobed 
dorsally  in  specimens  from  Leliefontein 
but  in  specimen  from  Wallekraal  expanded 
on  each  side  with  lobe  directed  laterally 
towards  ocular  sinus  and  another  directed 
dorsally);  underside  of  scape,  pedical  and 
proximal  flagellomeres;  continuous  anteri- 


Volume  16,  Number  2,  2007 


221 


or  band  on  pronotum  (narrowly  and 
pointedly  extended  a  little  along  dorso- 
lateral margin  and  broadly  continuous 
onto  humerus  and  beyond)  and  minute 
spot  on  postero-dorsal  angle  of  same;  small 
spot  at  top  of  mesopleuron  (specimen  from 
Wallekraal  only);  tegula  (except  for  median 
testaceaous  area);  scutellar  lamella  (other 
than  medially);  lower  two  thirds  of  meta- 
notum  (specimen  from  Wallekraal  only); 
minute  dots  dorsally  on  propodeum  (one 
specimen  from  Leliefontain  only)  or  small 
streak  unilaterally  on  angle  of  propodeum 
(specimen  from  Wallekraal  only);  narrow 
posterior  bands,  almost  reaching  sides,  on 
terga  I-VI.  The  following  are  light  reddish 
yellow:  mandible  (except  base  and  apex): 
labrum  (if  not  white);  posterior  bands, 
slightly  medially  expanded,  on  sterna; 
underside  of  trochanter  of  all  legs;  entire 
anterior  surface  of  fore  femur  as  well  as 
posterior  surface  of  basal  lamelliform  angle 
of  same;  underside  of  basal  half  of  mid 
femur  (most  specimens);  apices  of  femora 
and  entire  tibiae,  tarsi  and  claws  of  all  legs. 
Wing  membrane  sub-hyaline;  veins  brown. 

Length  5.8-6.0  mm.;  length  of  fore  wing 
3.8-3.9  mm.;  hamuli  circa  6. 

Head  in  front  view  1.33  X  as  wide  as 
long,  microreticulate,  moderately  shiny, 
with  shallow  punctures  (small  and  close 
on  frons,  slightly  larger  and  more  widely 
spaced  on  vertex).  POL:OOL  =  1:0.59. 
Clypeus  1.5  X  as  wide  as  long  in  midline; 
anterior  margin  widely  and  shallowly 
emarginate. 

Mesosoma  microreticulate,  moderately 
shiny,  with  punctures  larger  than  those 
on  head  (moderately  well  separated  on 
pronotum,  mesoscutum  and  scutellum). 

Tegula  short,  laterally  rounded,  with 
posterior  inner  corner  inwardly  produced. 

Fore  leg  much  modified;  coxa  and 
trochanter  enlarged;  femur  (Fig.  3)  greatly 
swollen,  its  posterior  surface  in  proximal 
half  markedly  concavely  excavate,  smooth 
and  very  shiny,  its  baso-ventral  region 
angulate  and  sublamellate;  first  tarsomere 
swollen,  excavate  and  setose  below;  second 


tarsomere  in  posterior  view  curved,  wide 
at  base  but  otherwise  narrow,  with  long  , 
backwardly  curved  setae;  middle  and  hind 
legs  beneath  with  trochanters  flattened  and 
with  femora  angulate,  flattened  in  proxi- 
mal half  and  longitudinally  grooved  in 
distal  half. 

Metasoma  moderately  shiny,  with  punc- 
tures finer  than  those  on  head.  Tergum  VII 
(Fig.  9)  drawn  out  apico-medially  into 
a  robust,  pointed,  dorsally  flattened  and 
apically  narrowly  rounded  process. 

Female:  Unknown,  none  of  the  specimens 
of  several  species  from  the  relevant  local- 
ities being  assignable  with  any  degree  of 
confidence  to  this  species. 

Etymology. — The  name,  namaquensis,  an 
adjective,  is  derived  from  the  Namaqua 
people  of  Namaqualand  and  refers  to  the 
provenance  of  the  species. 

Material  examined. — Holotype,  3,  SOUTH 
AFRICA:  NORTHERN  CAPE:  Leliefontein 
(30.23S  18.16E),  31.vii.2003  (C.  Mayer),  1  3 
(yellow  trap)  [AMG].  Paratypes:  SOUTH 
AFRICA:  NORTHERN  CAPE:  same  data  as 
holotype  but  date  15.ix.2003,  1  3  [AMG];  same 
data  as  holotype  but  date  22.viii.2004,  and  trap 
white,  1  3  [AMGJ;  W  of  Wallekraal  (30.21S 
17.26E),  8.X.1997  (F.  W.  and  S.  K.  Gess),  1  3  (ex 
nest  in  sand-filled  Trigonepmrus  shell)  [AMG]. 

Geographic  distribution. — Known  only 
from  two  localities  in  Namaqualand,  one 
in  the  coastal  sandveld,  the  other  in  the 
Kamiesberg.  In  the  former  locality  it  occurs 
together  with  Q.  conchicola  and  Q.  namaqua. 

Floral  associations. — Unknown. 

Nesting. — One  specimen,  freshly  eclosed 
and  with  wings  not  yet  fully  hardened, 
was  extracted  from  a  cell  of  a  nest  in  a  sand- 
filled  shell  of  the  desert  snail  Trigonephrus 
sp.  (Mollusca:  Gasteropoda:  Pulmonata: 
Dorcasiidae). 

Quartinia  obibensis  Gess,  new  species 

Diagnosis. — Large  to  very  large  (5.2- 
5.7  mm).  Fore  wing  with  Cula  and  2m-cu 
complete  and  as  thick  as  the  other  veins. 
Tegula  with  posterior  inner  corner  markedly 
inwardly  produced,  reddish  brown.   Both 


222  Journal  of  Hymenoptera  Research 

sexes  with  head  and  thorax  predominantly  Gaster   finely   microreticulate   but   shiny; 
black  with  limited  reddish-brown  markings;  punctures  finer  than  those  on  mesosoma, 
gaster  black  with  well  developed  reddish-  becoming  progressively  finer  distally. 
brown  posterior  bands  attaining  or  almost  Tegula  with  posterior  inner  corner  mark- 
attaining  side  of  terga.  Male  with  clypeus  edly  inwardly  produced, 
medially  depressed,  sparsely  punctured  and  Male:  Black.  The  reddish-brown  mark- 
shiny;  with  sternum   I  postero-medially  ings  as  in  the  female,  with  in  addition: 
raised  into  a  small  tubercle;  tubercle  viewed  labrum  (to  varying  degree);  in  some  speci- 
from  behind  transversely  subtriangular  with  mens  small  antero-lateral  spots  (occasion- 
a  narrowly  rounded  apex,  viewed  from  the  ally  joined  )  on  clypeus. 
side  sloping  gradually  anteriorly  and  falling  Length   5.2  mm;   length    of   fore   wing 
steeply  posteriorly  to  hind  margin  of  ster-  3.5  mm. 
num.  Tergum  VII  with  distinct  dorsal  and  POL:OOL  =  1:0.7 

lateral  surfaces;  apex  drawn  out  into  a  pair  of  Clypeus  medially  depressed,  non-micro- 
parallel  processes  flanking  narrow  and  slit-  sculptured  but  sparsely  punctured  and 
like  emargination;  emargination  produced  shiny.  Sternum  I  postero-medially  raised 
anteriorly  as  a  median  impression.  into  a  small  tubercle;  tubercle,  viewed  from 
Description. — Female:  Black.  The  follow-  behind,  transversely  subtriangular  with 
ing  are  reddish-brown:  mandibles  (except  a  narrowly  rounded  apex,  anteriorly  grad- 
base);  underside  of  pedicel  and  flagello-  ually  sloping,  posteriorly  falling  steeply  to 
meres;  anterior  margin  of  pronotum  and  hind  margin  of  sternum.  Tergum  VII 
postero-dorsal  angle  of  same;  tegula;  cres-  (Fig.  10)  with  distinct  dorsal  and  lateral 
cent  (in  some  specimens  broken  up  into  surfaces;  apex  drawn  out  into  a  pair  of 
spots)  posteriorly  and  laterally  on  disk  of  parallel  processes  flanking  narrow  and  slit- 
scutellum;  scutellar  lamella;  in  some  speci-  like  emargination;  emargination  produced 
mens  lower  half  of  metanotum;  posterior  anteriorly  as  a  median  impression, 
bands  attaining  or  almost  attaining  sides  Etymology. — The  name,  obibensis,  an  ad- 
on  terga  I-V;  that  on  I  of  even  width  and  jective,  is  derived  from  the  Obib  Mountains 
covering  about  half  of  tergum;  those  of  in  the  Sperrgebiet  of  south-western  Nami- 
terga  I  I-V  progressively  narrower,  undu-  bia,  the  site  from  which  the  largest  number 
late,  expanded  medially  and  laterally  and  of  specimens  was  obtained. 

attaining  or  almost  attaining  sides  of  terga); 

•     i         *         «.         ™  t\7    „„,w  n(  c°  I  Material  examined.— Holotype:  $,  NAMIBIA: 

apical  spot  on  tergum  IV    apex  of  femur,  „                    _,  ,                  yr     "'        „,„„_. 

*\          *\                              ,            l  i     t.\     £    ii  Sperrgebiet,   Obib   camp    site     28.00S    16.39E ), 

entire  tibia,  tarsomeres  (except  last)  of  all  ,\  .    ®n   '     TA7            *   v    _ v    .   . 

,      ,            .  14.ix.2003  (F.  W.  and  S.  K.  Gess)  (ex  nest  in 

legs.    Last    tarsomere    and    claws   brown.  sand.filled  Trigonephrus  shell/  emerged  in  the 

Wings  slightly  darkened;  veins  brown.  lab  15_22.x.2003)  [AMG].  Paratypes:  NAMIBIA: 

Length    5.2-5.7  mm    (average    of   4:5.4  Sperrgebiet,   W   of   Klinghardtberge   (27.17S 

mm);    length    of    fore    wing    3.6-3.7  mm  15.36E),  20.ix.2003  (F.  W.  and  S.  K.  Gess),  4  99 

(average  of  4:3.7  mm);  hamuli  5.  (ex   nests   in   sand-filled   Trigonephrus   shells) 

Head  in  front  view  1.35  X  as  wide  as  long,  [AMG];   Sperrgebiet,   Klinghardtberge   (27.19S 

microreticulate,  matt,  with  inconspicuous,  15.46E),  10.ix.2005  (F.  W.  and  S.  K.  Gess),  1  9 

very  shallow,  fine  punctures.  POL:OOL  =  (reared  ex  Quartinia  nest  in  sand-filled  Trigone- 

1:0.7.  Clypeus  1.33  X  as  wide  as  long  (length  Phrus   shell)   tAMGl;   Sperrgebiet,   Klinghardt- 

_            J\.     u   „           c              •      «        1  o  n/  berge  (27.30S,  15.44E),  10.ix.2005  (F.  W.  and  S. 

measured  to  bottom  of  emargination;  1.2  X  T,  %   \   «     '              '             «        .  . 

.,                    j         1        1      r                 t    ,        1  K.  Gess),  1  9,  1(5  (reared  ex  Quart  una  nests  in 

it  measured  to  level  of  antero-lateral  an-  ,  c.u    ,  ^  .          ,          ,    ,,  .  taa*/-!    a 

sand-filled  Trigonephrus  shells)  [AMG];  Aus  - 

gles);  anterior  margin  smooth,  shiny,  shal-  Rosh  pinah  (27  44S  16  42E)    25  ix  2003  (F.  w. 

lowly  and  evenly  emarginate.  and  s   K   Gess)^  1  s  (visiting  white  centered, 

Mesosoma    microreticulate    with   close,  purplish-pink  rayed  Drosanthemum  spv  Aizoa- 

shallow,    fine    punctures,    slightly    shiny,  ceae:  Mesembrianthema)  [AMG;  ];  Sperrgebiet, 


Volume  16,  Number  2,  2007 


223 


between  Aurusberg  and  Scorpion  Mine  (27.45S 
16.32E),  15.ix.2003  (F.  W.  and  S.  K.  Gess),  1  9,  2 
SS  (ex  nests  in  sand-filled  Trigonephrus  shells) 
[AMG];  Sperrgebiet,  Scorpion  Mine  (27.49S 
16.35E),  15.ix.2003  (F.  W.  and  S,.  K.  Gess),  4  99, 

1  j  (ex  nests  in  sand-filled  Trigonephrus  shells) 
[AMG];  Sperrgebiet,  Obib  camp  site  (28.00S 
16.39E),  14.ix.2003  (F.  W.  and  S.  K.  Gess),  37 
99,  13  $<$  (ex  nests  in  sand-filled  Trigonephrus 
shells;  26  99,  2  J  J  emerged  in  the  lab.  15- 
22.X.2003;  7  99,  2  S3  emerged  in  lab.  at  a  later 
date  )  [AMG];  12.8  km  S  Rosh  Pinah  (28.03S 
16.51E)  ll.ix.1996  (F.  W.,  S.  K.  and  R.  W.  Gess),  3 
99/  3  0*6*  (ex  nests  in  sand-filled  Trigonephrus 
shells;  2  99,  3  $$  emerged  in  lab.)  [AMG]. 
SOUTH  AFRICA:  NORTHERN  CAPE:  Richters- 
veld,  W  of  Brandkaros  (28.29S  16.40  E), 
15.ix.1996  (F.  W.,  S.  K.  and  R.  W.  Gess),  3  99,  1 
6*  (ex  nests  in  sand-filled  Trigonephrus  shells) 
[AMG].  WESTERN  CAPE:  Yzerfontein  (33.20S 
18.10E),  15.X.2006  (D.  W.,  G.  T.  and  G.  M.  Gess), 

2  $$  (ex  Theba  pisana  shells)  [AMG]. 

Geographic  distribution. — Q.  obibensis  is 
known  from  Namibia,  from  a  limited  area 
in  the  southern  half  of  the  Desert  and 
Succulent  Steppe  (Winter  Rainfall  Area)  of 
Giess  (1971),  from  South  Africa  from 
a  nearby  locality  in  the  Richtersveld  and 
from  a  coastal  site  in  the  Western  Cape.  In 
the  north  of  its  range  it  occurs  together 
with  Q.  conchicola,  Q.  rufigicola  and  Q. 
vexillata  and  in  the  south  with  Q.  bonaespei 
and  Q.  namaqua. 

Floral  associations. — Aizoaceae:  Mesem- 
bryanthema  (Drosanthemum). 

Nesting. — Throughout  its  presently 
known  distributional  area  found  nesting 
most  commonly  in  sand-filled  shells  of  the 
desert  snail  Trigonephrus  sp.  (Mollusca: 
Gasteropoda:  Pulmonata:  Dorcasiidae).  At 
one  coastal  locality  in  the  Western  Cape 
obtained  from  shells  of  the  exotic  Theba 
pisana  (Mull.)  (Mollusca:  Gasteropoda: 
Pulmonata:  Helicidae).  For  further  details 
see  Gess  and  Gess  (1999). 

Discussion. — See  under  Q.  namaqua. 

Quartinia  refugicola  Gess,  new  species 

Quartinia  sp.  (smaller  sp.)  (Gess  and  Gess  1999, 
nesting;  Greathead  1999,  2006,  bombyliid 
parasite). 


Diagnosis. — Medium  sized  to  large  (4.1- 
5.2  mm  long).  Fore  wing  with  Cula  and 
Im-cu  complete  and  as  thick  as  other  veins. 
Tegula  with  posterior  inner  corner  inward- 
ly produced.  Posterior  bands  on  terga 
reaching  lateral  margins. 

Description. — Female:  Black.  The  follow- 
ing are  yellow  or  yellow  merging  into 
brownish  yellow:  underside  of  flagello- 
meres;  short,  narrow,  transverse  band  (in 
some  specimens  reduced  to  pair  of  small 
marks,  in  others  totally  absent)  medially  on 
pronotum  and  in  some  specimens  a  minute 
dot  on  postero-dorsal  angle  of  same;  tegula 
(except  for  testaceous  medial  spot);  nar- 
row, medially  interrupted,  lamellate  mar- 
gin of  scutellum;  in  some  specimens 
median  part  of  metanotum;  narrow  poste- 
rior bands  reaching  lateral  margins  on 
terga  I-V  (that  of  tergum  I  widest,  others 
progressively  narrower);  in  some  speci- 
mens a  diffuse  posterior  band  on  sternum 
II;  extreme  apex  of  femur,  entire  tibia 
(except  for  elongate  dark  mark  on  posterior 
surface)  and  tarsomeres  of  all  legs  (except 
in  some  specimens  brown  terminal  tar- 
someres of  middle  and  hind  legs).  Mandi- 
ble with  distal  half  bright  ferruginous; 
labrum  brown.  Wings  subhyaline;  veins 
brown. 

Length  4.5-5.2  mm  (average  of  5:4.8 
mm);  length  of  fore  wing  3.0-3.4  mm 
(average  of  5:3.2  mm);  hamuli  5-6. 

Head  in  front  view  1.3  X  as  wide  as  long; 
clypeus  1.5  X  as  wide  as  long  (length 
measured  to  bottom  of  emargination); 
POL:OOL  =  1:0.6.  Clypeus  very  closely 
microsculptured,  with  barely  discernable 
shallow  punctures;  Irons  and  vertex  simi- 
larly microsculptured  but  more  obviously 
punctured  (especially  in  region  of  ocelli); 
mesosoma  microsculptured  with  obvious 
shallow  punctures  slightly  larger  than 
those  on  vertex  and  with  interstices  of 
puncture  width  or  less;  gaster  closely  and 
finely  punctured. 

Male:  Black.  Pale  markings  as  in  female 
but  with  the  addition  of:  in  some  speci- 
mens sub-basal  spot  on  mandible  between 


224 


Journal  of  Hymenoptera  Research 


black  base  and  ferruginous  distal  half; 
in  some  specimens  part  of  the  clypeus 
(ranging  in  extent  from  pair  of  antero- 
lateral spots,  to  uninterrupted  anterior 
margin,  to  most  of  disc  with  exception  of 
region  below  antennal  sockets);  in  all 
specimens  narrow  posterior  band  on  ter- 
gum  VI  and  in  most  specimens  apices  of 
tergum  VII. 

Length  4.1-4.3  mm  (average  of  5:4.2 
mm);  length  of  fore  wing  2.8-2.9  mm 
(average  of  5:2.8  mm);  hamuli  4. 

Head  in  front  view  1.37  X  as  wide  as 
long;  clypeus  convex,  1.5  X  as  wide  as 
long;  POL:OOL  =  1:0.6.  Microsculpture 
and  puncturation  as  in  female.  Tergum  VII 
(Fig.  11)  dorsally  slightly  depressed  (flat- 
tened) and  its  apical  margin  with  a  narrow 
V-shaped  median  emargination  flanked  by 
narrowly  rounded  projections. 

Etymology. — The  name  refugicola  is  a  com- 
pound word  formed  from  the  Latin  words 
refugium  -  ii  (n)  -  a  place  of  refuge,  and  cola 
-  a  dweller.  It  serves  to  draw  attention  to 
the  species'  association  with  sand-filled 
cavities  in  which  its  nests  are  sheltered 
from  prevailing  winds. 

Material  examined. — Holotype:  6\  NAMIBIA: 
12.8  km  S  of  Rosh  Pinah  (28.03  S  16.51E), 
ll.ix.1996  (F.  W.,  S.  K.  and  R.  W.  Gess)  (on 
ground)  [AMG].  Paratypes:  NAMIBIA:  Aus 
(26.39S  16.15E),  25.viii.2002  (F.  W.  and  S.  K. 
Gess),  1  9  (visiting  yellow  flowers  of  Leysera 
tenella  DC,  Asteraceae)  [AMG];  Sperrgebiet, 
Kaukausib  Spring  -  Grillental  (26.58S  15.31E), 
5.ix.2002  (F.  W.  and  S.  K.  Gess),  4  99  (visiting 
white  flowers  of  Zygaphyllum  clavatum  Schltr.  & 
Diels,  Zygophyllaceae)  [AMG];  Sperrgebiet, 
near  Grillental  (26.59S  15.23E),  5.ix.2002  (F.  W. 
and  S.  K.Gess),  1  6*  (visiting  yellow  flowers  of 
Foveolina  albida  (DC.)  Kallersjo,  Asteraceae) 
[AMG];  Sperrgebiet,  Grillental  (27.00S  15.21E), 
8.ix.2005  (F.  W.  and  S.  K.  Gess),  3  99,  1  $ 
(visiting  white  flowers  of  Zygophyllum  sp.) 
[AMG];  Sperrgebiet,  Klinghardtberge,  Tsabiams 
Camp  (27.10  S  15.42E),  4.ix.2002  (F.  W.  and  S.  K. 
Gess),  1  9  (visiting  yellow  flowers  of  Dimor- 
photheca  polyptera  DC,  Asteraceae)  [AMG]; 
Sperrgebiet,  Klinghardtberge  (27.14S  15.43E), 
l-3.ix.2002  (F.  W.  and  S.  K.  Gess),  5  99  (1  9 


visiting  flowers  of  ?  Cotula  sp.,  Asteraceae;  1  9 
visiting  yellow  flowers  of  Pteronia  sp.,  Aster- 
aceae; 3  99  visiting  yellow  flowers  of  Zygophyl- 
lum simplex  L.,  Zygophyllaceae)  [AMG];  Sperr- 
gebiet, Klinghardtberge  (27.14S  15.44E), 
2.ix.2002  (F.  W.  and  S.  K.  Gess),  6  99,  1  J  (3  99 
visiting  yellow  flowers  of  Pteronia  sp.,  Aster- 
aceae; 3  99  visiting  apricot  coloured  flowers  of 
Phyllobolus  occulatus  (N.E.Br.)  Gerbaulet,  Aizoa- 
ceae:  Mesembryanthema;  1  6"  visiting  pink 
flowers  of  Sarcocaulon  sp.,  Geraniaceae)  [AMG]; 
Sperrgebiet,  W  of  Klinghardtberge  (27.17S 
15.36E),  20.ix.2003  (F.  W.  and  S.  K.  Gess),  5  99, 
3  S3  (ex  nests  in  sand-filled  Trigonephrus  shells) 
[AMG];  Sperrgebiet,  Klinghardtberge  (27.18S 
15.54E),  2.ix.2002)  (F.  W.  and  S.  K.  Gess),  1  6" 
[AMG];  Sperrgebiet,  Klinghardtberge  (27.19S 
15.46E),  10.ix.2005  (F.  W.  and  S.  K.  Gess),  2  99, 
1  S  (reared  from  larvae  ex  Quartinia  nests  ex 
sand-filled  Trigonephrus  shells)  [AMG];  Sperrge- 
biet, Klinghardtberge,  Nomitsas  (27.27S  15.52E), 
31.viii.20Q2  (F.  W.  and  S.  K.  Gess),  1  9,  1  o  (ex 
sand-filled  Trigonephrus  shells)  [AMG];  Sperrge- 
biet, Uguchab  River,  NW  of  Aurus  Mountains 
(27.31S  16.12E),  17.ix.2003  (F.  W.  and  S.  K.  Gess), 
31  99,  20  (J6*  (17  99,  10  26*  ex  sand-filled 
Trigonephrus  shells;  14  99,  10  $<$  ex  nests  in 
sand-filled  Trigonephrus  shells)  [AMG];  12.8  km  S 
of  Rosh  Pinah  (28.03  S  16.51E),  ll.ix.1996  (F.  W., 
S.  K.  and  R.  W.  Gess),  29  99,  32  6*6"  (21  99,  26  SS 
on  ground;  1  9  visiting  yellow  flowers  of  Cotula 
sp.,  Asteraceae;  1  9  on  blue  rayed  Filicia  sp., 
Asteraceae;  1  9,  1  j  visiting  yellow  flowers  of 
Hirpicium  sp.,  Asteraceae;  1  9, 1  6*  visiting  yellow 
flowers  of  Osteospermum  sp.,  Asteraceae;  1  9 
visiting  yellow  flowers  of  Grielum  sp.,  Neurada- 
ceae;  3  99,  4  ^J  ex  nests  in  sand-filled  Trigone- 
phrus shells;  2  SS  ex  sand-filled  cavities  in 
calcrete)  [AMG];  Sperrgebiet,  W  of  Obib  Moun- 
tains (28.08S  16.42E),  15.ix.2003  (F.  W.  and  S.  K. 
Gess),  1  9  (ex  nest  in  sand-filled  Trigonephrus 
shell)  [AMG];  E  of  Oranjemund,  28  km  from 
checkpoint  on  road  to  Sendelingsdrif  (28.26S 
16.42E),  25.ix.1997  (F.  W.  and  S.  K.  Gess),  1<J  (ex 
nest  in  sand-filled  Trigonephrus  shell)  [AMG];  E 
of  Oranjemund  (28.30S  16.36E),  22.ix.1997  (F.  W. 
and  S.  K.  Gess),  1  6  (ex  nest  in  sand-filled 
Trigonephrus  shell)  [AMG].  SOUTH  AFRICA: 
NORTHERN  CAPE:  W  of  Brandkaros  (28.29S 
16.40E),  13-15.ix.1996  (F.  W.,  S.  K.  and  R.  W. 
Gess),  4  99  (1  dead),  1  6  (dead)  (ex  nests  in  sand- 
filled  Trigonephrus  shells)  [AMG];  60  km  N  of 
Port  Nolloth  (28.47S  16.38E),  27.ix.1997  (F.  W. 


Volume  16,  Number  2,  2007 


225 


and  S.  K.  Gess),  1  $  (ex  nest  in  sand-filled 
Trigonephrus  shell)  [AMG]. 

Geographic  distribution. — Quartinia  refugi- 
cola  is  known  from  Namibia,  from  numerous 
localities  in  the  Desert  and  Succulent  Steppe 
(Winter  Rainfall  Region)  of  Giess  (1971)  and 
from  the  immediately  adjacent  area  across 
the  Orange  River  in  the  Northern  Cape  of 
South  Africa.  It  occurs  together  with  Q. 
conchicola,  Q.  obibensis  and  Q.  vexillata. 

Floral  associations. — Known  in  associa- 
tion with  Aizoaceae:  Mesembryanthema 
(Pln/llobolus),  Asteraceae  (Cotula,  Dimor- 
phothcca,  Filicia,  Foveolina,  Hirpiciwn,  Ley- 
sera,  Osteospermum  and  Pteronia),  Gerania- 
ceae  (Sarcocaulon),  Neuradaceae  (Grielum) 
and  Zygophyllaceae  (Zygophyllum). 

Nesting. — Throughout  its  presently 
known  distributional  area  most  commonly 
found  nesting  in  sand-filled  shells  of  the 
desert  snail  Trigonephrus  sp.  (Mollusca: 
Gasteropoda:  Pulmonata:  Dorcasiidae), 
less  commonly  in  sand-filled  cavities  in 
calcrete  rocks.  See  also  Gess  and  Gess 
(1999).  At  several  localities  in  the  Sperrge- 
biet  nests  have  been  found  to  be  parasitised 
by  Apolysis  hesseana  Evenhuis  and  Great- 
head  (Bombyliidae:  Usiinae:  Apolysini). 
See  also  Greathead  (1999:155;  2006:  5). 

Quartinia  vexillata  Gess,  new  species 

Diagnosis. — Large  to  very  large  (5.2- 
6.5  mm).  Fore  wing  with  Cula  and  2>n-cu 
complete  and  as  thick  as  other  veins.  Both 
sexes  with  vertex  behind  posterior  ocelli 
evenly  convex;  with  fore  coxa  swollen 
basally  and  anteriorly  produced,  very 
markedly  so  in  male,  less  so  in  female 
where  swelling,  however,  forms  a  rounded 
right  angle.  Male  with  fore  femur  greatly 
enlarged,  robust,  proximally  produced 
ventrally  to  form  a  sturdy,  subquadrate 
flange,  distally  markedly  downcurved; 
flange  with  its  posteriorly  facing  surface 
markedly  concave  with  pronounced  distal 
angles  and  its  anteriorly  facing  surface 
convex  with  a  pronounced  submedian 
distal  tubercle;  tibia  robust  with  dense 
setae  on  lower  surface. 


Description. — Female:  In  general  facies 
similar  to  male  (described  below)  but 
with  legs  and  last  tergum  simple.  Head 
without  any  pale  markings.  Specimens 
from  between  Alexander  Bay  and  Port 
Nolloth  have  the  mesosoma  and  gaster 
with  both  yellowish  white  and  reddish 
brown  markings  very  similar  to  those  of 
males  from  the  same  population;  speci- 
mens from  SSE  of  Grillental  and  from  Obib 
have  the  markings  on  the  mesosoma 
tending  to  reddish  brown.  Microsculpture 
(shagreening)  of  head  and  mesosoma 
somewhat  coarser  and  these  parts,  in 
particular  mesoscutum,  semi-matt  rather 
than  shiny. 

Length  5.8-6.5  mm  (average  of  7  = 
6.1  mm);  length  of  fore  wing  3.9-4.4  mm 
(average  of  7  =  4.1  mm);  hamuli  6. 

Head  broad,  1.33  X  as  wide  as  long; 
POL:OOL  =  1:0.5.  Vertex  behind  posterior 
ocelli  evenly  convex.  Fore  coxa  enlarged, 
basally  markedly  and  roundly  anteriorly 
produced. 

Male:  Head  and  mesosoma  black,  gaster 
and  greater  part  of  femur  of  all  legs  very 
dark  brown  to  almost  black.  The  following 
are  yellowish  white:  lower  aspect  of  scape 
(excluding  radicle)  and  pedicel;  labrum  (in 
one  specimen  testaceous);  clypeus  (other 
than  for  irregular  area  below  antennal 
socket);  a  small  transverse  spot  situated 
on  either  side  of  midline  of  frons  immedi- 
ately above  frontoclypeal  suture  (in  one 
specimen  only);  narrow  paraocular  streak 
from  mandibular  insertion  to  level  of  top  of 
antennal  socket;  short  (in  one  specimen 
almost  medially  interrupted)  and  laterally 
widening  transverse  band  on  dorsum  of 
pronotum  and  minute  spot  at  postero- 
dorsal  angle  of  same;  humeral  streak  of 
varying  length;  anterior  and  posterior 
thirds  of  tegula  (median  third  clear,  testa- 
ceous); medially  interrupted  band  on  la- 
mellate margin  of  scutellum;  distal  portion 
of  flange  on  fore  femur.  The  following  are 
various  shades  of  light  reddish-brown: 
mandible  (other  than  base);  flagellomeres 
(other  than  for  dark  suffusion  on  upper 


226 


Journal  of  Hymenoptera  Research 


surface);  posterior  bands  (slightly  widened 
medially,  narrowed  laterally,  and  not  quite 
attaining  lateral  margins  of  terga)  on  terga 
I-VI;  streak  on  anteriorly  protruding  por- 
tion of  fore  coxa  (in  one  specimen)  and 
lower  surface  of  middle  and  hind  coxae; 
all  trochanters;  distal  portion  (up  to 
almost  half  the  length)  of  anterior 
aspect  of  fore  femur;  basal  flattened  lower 
surface  of  middle  femur;  apex  of  middle 
and  hind  femora;  tibia  and  tarsus  of 
all  legs.  Venation  light  brown  at  base  of 
wings,  otherwise  very  dark  brown.  Wing 
membrane  very  slightly  browned,  a  little 
darker  on  fore  wing  in  and  beyond 
marginal  cell. 

Length  5.2-6.4  mm;  length  of  fore  wing 
3.6-3.8  mm;  hamuli  6. 

Head,  mesosoma  and  terga  I-VI  very 
finely  microsculptured  (shagreened)  but 
nevertheless  shiny,  with  moderately  sized 
punctures;  punctures  on  head  and  terga 
somewhat  shallow  and  undefined  with 
interspaces  generally  less  than  puncture 
diameter,  those  on  mesosoma  deeper  and 
well  defined  with  interspaces  at  least  on 
mesoscutum  often  greater  than  puncture 
diameter.  Tergum  VII  without  microsculp- 
ture;  punctures  more  pronounced  than 
those  on  other  terga,  irregularly  spaced, 
some  separated  by  wide  interspaces  and 
others  coalescing. 

Setation  on  head  and  particularly  on 
body  sparse  and  short  throughout,  more 
noticeable  on  tibiae  and  tarsi  and  strikingly 
developed  on  underside  of  front  tibia 
where  dense  and  long. 

Head  broad,  1.45  X  as  wide  as  long; 
POL:OOL  =  1:0.5.  Vertex  behind  posterior 
ocelli  evenly  convex; 

Tegula  with  posterior  inner  corner  in- 
wardly produced.  Wing  venation  with 
Cula  and  2m-cu  complete  and  as  thick  as 
other  veins. 

Fore  leg  uniquely  and  greatly  modified; 
coxa  enlarged,  basally  markedly  and 
roundly  anteriorly  produced;  femur 
(Fig.  4)  greatly  enlarged,  robust,  proximal- 
ly  produced  ventrally  to  form  a  sturdy, 


subquadrate  flange,  distally  markedly 
downcurved;  flange  with  its  posteriorly 
facing  surface  markedly  concave  with  pro- 
nounced distal  angles  and  its  anteriorly 
facing  surface  convex  with  a  pronounced 
submedian  distal  tubercle;  tibia  robust 
with  dense  setae  on  lower  surface. 

Middle  and  hind  femora  robust,  mark- 
edly angled  below  and  with  lower  surface 
both  proximal  and  distal  to  angle  distinctly 
flattened  (more  so  on  middle  than  on  hind 
femur). 

Tergum  VII  (Fig.  12)  in  posterior  half 
with  dorsal  surface  raised  laterally  and 
slightly  concave  medially,  produced  api- 
cally  and  with  a  deep,  narrow,  slightly  sub- 
parallel-sided  median  slit. 

Etymology. — The  name  vexillata  is  formed 
from  the  Latin  noun  vexillum  meaning 
a  flag  or  standard.  It  refers  to  the  conspic- 
uously modified  front  femur  of  the  male 
which  may  possibly  have  a  communicatory 
role  in  courtship  behaviour. 

Material  examined.— Holotype:  &  SOUTH 
AFRICA:  NORTHERN  CAPE:  23  km  S  of 
Alexander  Bay  (28.46S  16.37E),  11.x.  2000  (F. 
W  and  S.  K.  Gess)  (on  ground)  [AMG].  Para- 
types:  NAMIBIA:  Sperrgebiet,  main  north/ 
south  road,  47  km  SSE  of  Grillental  (27.23S 
15.32E),  6.ix.2002  (F.  W.  and  S.  K.  Gess),  2  99  (on 
ground  next  to  Drosanthemum  sp.,  Aizoaceae: 
Mesembryanthema)  [AMG];  Sperrgebiet,  Obib 
camp  site  (28.00S  16.39E),  14.ix.2003  (F.  W.  and 
S.  K.  Gess),  9  99  (7  99  visiting  yellow  flowers  of 
Cephalophyllum  sp.,  Aizoaceae:  Mesem- 
bryanthema; 2  99  visiting  yellow  flowers  of 
Othoiuia  cylindrica  (Lam.)  DC,  Asteraceae) 
[AMG].  SOUTH  AFRICA:  NORTHERN 
CAPE:  23  km  S  of  Alexander  Bay  (28.46S 
16.37E),  11.x.  2000  (F.  W  and  S.  K.  Gess),  5  99 
(4  99  visiting  pink  flowers  of  Drosanthemum  sp., 
Aizoaceae:  Mesembryanthema;  1  9  visiting 
purple-centred  white  flowers,  Aizoaceae:  Me- 
sembryanthema) [AMG];  60  km  N  of  Port 
Nolloth  (28.47S  16.38E),  27.  ix.  1997)  F.  W.  and 
S.  K.  Gess),  6  99,  1  $  (2  99,  £  visiting  pale  pink 
flowers  of  Drosanthemum  sp.;  4  99  on  ground) 
[AMG]. 

Geographic  distribution. — The  species  is 
known    from   Namibia    from    the    Desert 


Volume  16,  Number  2,  2007 


227 


and  Succulent  Steppe  (Winter  rainfall  area)  Floral   associations. — Aizoaceae:   Mesem- 

of  Giess  (1971)  and  from  South  Africa  from  bryanthema    {Cephalophyllum,    Drosanthe- 

the  adjoining  northern  Strandveld  of  the  mum);  Asteraceae  (Othonna). 

West  Coast   of  Acocks   (1953).    It   occurs  Nesting. — Unknown;   probably   utilizing 

variously  together  with  Q.  conchicola,  Q.  sand-filled  snail  shells  as  a  nesting  niche. 

obibcnsis  and  Q.  refugicola.  Discussion. — See  under  Q.  conchicola. 


KEY  TO  SPECIES  NESTING  IN  SAND-FILLED  SNAIL  SHELLS  OR  (VEXILLATA) 

PRESUMED  TO  DO  SO 
Males 


4. 
5. 


6. 


Fore  legs  not  modified 2 

Fore  legs  markedly  modified    5 

Sternum  I  not  modified    3 

Sternum  I  posteriorly  raised  into  a  tubercle 4 

Large  (5.0-5.4  mm  long);  black  with  white  markings;  tegula  with  white  anterior  and 
posterior  markings  contrasting  markedly  with  dark  brown  to  black  median  part; 
pale  posterior  bands  on  terga  not  extending  onto  sides;  clypeus  and  labrum 

white australis  Gess  n.  sp. 

Medium  (4.1-4.3  mm  long);  black  with  yellow  to  brownish-yellow  markings;  tegula 
with  pale  anterior  and  posterior  markings  not  contrasting  markedly  with 
testaceous  median  part;  pale  posterior  bands  on  terga  reaching  lateral  margins 

refugicola  Gess  n.  sp. 

Clypeus  convex    namaqua  Gess  n.  sp. 

Clypeus  depressed  to  concave    obibensis  Gess  n.  sp. 

Posterior  bands  on  terga  white;  not  contrasting  in  colour  with  markings  on  head  and 
mesosoma;  fore  femur  (Fig.  3)  greatly  swollen,  its  posterior  surface  in  proximal  half 
markedly  concavely  excavate,  smooth  and  very  shiny,  its  baso-ventral  region 

angulate  and  sublamellate namaquensis  Gess  n.  sp. 

Posterior  bands  on  terga  reddish-brown  or  bright  reddish-orange,  in  most  specimens 
contrasting  in  colour  with  markings  on  head  and  mesosoma;  fore  femur  differently 

formed     6 

Tibiae  and  tarsi  of  all  legs  predominantly  black;  fore  femur  (Fig.  1)  greatly  swollen, 
postero-basally  with  a  pointed  tubercle,  its  posterior  surface  depressed,  smooth  and 

very  shiny  and  forming  an  angle  with  ventral  surface     bonaespei  Gess  sp.  n. 

Tibiae  and  tarsi  of  all  legs  predominantly  light  reddish-brown    7 

Vertex  behind  posterior  ocelli  evenly  convex;  fore  coxa  swollen  basally  and  markedly 
anteriorly  produced;  fore  femur  (Fig.  4)  greatly  enlarged,  robust,  proximally 
produced  ventrally  to  form  a  sturdy  subquadrate  flange,  distally  markedly 
downcurved;  flange  with  its  posterior  facing  surface  markedly  concave  with 
pronounced  distal  angles  and  its  anterior  facing  surface  convex  with  a  pronounced 

submedian  distal  tubercle    vexillata  Gess  n.  sp. 

Vertex  behind  posterior  ocelli  depressed,  somewhat  concave;  fore  coxae  unmodified; 
fore  femur  (Fig.  2)  enlarged,  excavated  beneath  and  undulate  postero-ventmlly 
conchicola  Gess  n.  sp. 


Females 

Species  not  included:  namaquensis  Gess  n.  sp.  (9  not  known) 


1. 


Markings  on  mesosoma  and  gaster  generally  concolorous 2 


228 


Journal  of  Hymenoptera  Research 


Markings  on  mesosoma  and  gaster  not  of  same  colour;  posterior  bands  on  terga 
reddish-brown  or  bright  reddish-orange,  generally  contrasting  with  pale  markings 
on  mesosoma     5 

2.  Black  with  white  markings;  tegula  with  anterior  and  posterior  markings  contrasting 

markedly  with  dark  brown  to  black  median  part;  posterior  bands  on  terga  not 

extending  onto  sides    anstralis  Gess  sp.  n. 

Black  with  yellow,  brownish-yellow  or  reddish-brown  markings;  tegula  with  anterior 
and  posterior  markings  not  contrasting  markedly  with  median  part;  posterior 
bands  on  terga  variously  developed    3 

3.  Posterior  bands  on  terga  reaching  lateral  margins;  scutellar  disk  black  and  scutellar 

lamella  yellow  to  brownish  yellow    refugicola  Gess  sp.n. 

Without  this  combination  of  characters    4 

4.  Mesoscutum    and    scutellum   with    interstices   between   punctures    not   obviously 

microreticulate  (shagreened);  scutellar  disk  and  scutellar  lamella  black;  scutellar 
lamella  at  most  slightly  flattened  postero-medially;  metanotum  not  transversly 

impressed,  black  throughout namaqua  Gess  n.  sp. 

Mesoscutum  and  scutellum  with  interstices  between  punctures  very  obviously 
microreticulate  (shagreened);  scutellar  disk  laterally  and  medially  with  reddish- 
brown  markings  and  scutellar  lamella  of  same  colour;  scutellar  lamella  slightly 
emarginate   postero-medially;    metanotum    transversely    impressed    with   lower 

section  reddish  brown  and  contrasting  with  almost  black  upper  section 

obibensis  Gess  n.  sp. 

5.  Tibiae  and  tarsi  of  all  legs  predominantly  black bonaespei  Gess  sp.  n. 

Tibiae  and  tarsi  of  all  legs  predominantly  light  reddish-brown    6 

6.  Vertex  behind  posterior  ocelli  evenly  convex;  fore  coxae  swollen  basally  and  markedly 

anteriorly  produced vcxillata  Gess  n.  sp. 

Vertex  behind  posterior  ocelli  depressed,  somewhat  concave;  fore  coxae  unmodi- 
fied         conchicola  Gess  n.  sp. 


B)     Other  species 

Quartinia  femorata  Gess,  new  species 

Diagnosis. — Very  large  to  gigantic  (5.8- 
6.7  mm  long).  Fore  wing  with  Cula  and 
2m-cu  complete  and  as  thick  as  the  other 
veins.  Tegula  with  posterior  inner  corner 
inwardly  produced.  Both  sexes  predomi- 
nantly yellow.  Male  fore  femur  robust, 
notched  ventrally  in  basal  third  and  with 
a  distally  directed,  apically  rounded,  la- 
mellate process. 

Description. — Female:  Predominantly  yel- 
low. Black  greatly  reduced  leaving  only: 
occiput;  irregular  median  band  on  vertex 
(posteriorly  wide  along  occipital  carina  but 
anteriorly  narrowed  and  closely  encom- 
passing ocelli)  and  on  frons  (on  upper  half 
of  similar  width  to  part  encompassing 
ocelli  but  on  lower  half  trifid  with  middle 


arm  and  outcurved  lateral  arms  reaching 
clypeal  suture  and  antennal  sockets  re- 
spectively); propleuron  (in  greater  part) 
and  presternum;  median  and  parapsidal 
bands  on  mesoscutum  (median  band  wide 
at  anterior  margin,  narrowing  posteriorly; 
parapsidal  bands  not  reaching  anterior 
margin  and  of  even  width  throughout); 
small  antero-median  mark  on  scutellum; 
anterior  half  of  propodeal  dorsum  and 
small  spot  on  each  side  at  bottom  of 
propodeal  declivity;  transverse  marking 
(either  continuous  or  broken  up  into  three) 
on  declivity  of  tergum  I;  abbreviated 
anterior  transverse  bands  (only  visible  if 
metasoma  is  downwardly  flexed)  on  terga 
II  and  III.  The  following  are  various  shades 
of  light  reddish-brown:  mandibular  teeth; 
antennal  club  (apex  of  last  flagellomere 
dark  brown);  last  one  or  two  tarsomeres 


Volume  16,  Number  2,  2007 


229 


(arolia  dark  brown);  usually  concealed 
anterior  third  of  terga  II— VI  and  poorly 
defined  laterally  abbreviated  and  medially 
interrupted  pre-apical  transverse  bands  on 
terga  II- V.  Tegulae  yellowish-white  except 
for  unpigmented  translucent  central  area 
and  outer  margin.  Wing  membrane  hya- 
line; costa,  subcosta,  media,  thickening  at 
junction  of  Rs  &  M,  parastigma  and  stigma 
light  brown,  rest  of  venation  contrastingly 
dark  brown. 

Length  5.8-6.7  mm  (average  of  6:6.3  mm; 
length  of  front  wing  3.8-4.3  mm  (average 
of  6:4.1  mm);  hamuli  7. 

Head,  thorax  and  gaster  sparsely  cov- 
ered with  short,  semi-erect  pale  pilosity, 
slightly  longer  and  most  noticeable  on 
head,  declivity  of  propodeum,  declivity  of 
tergum  I,  and  sternum  VI. 

Head  in  front  view  1.25  X  as  wide  as 
long,  microreticulate,  with  close,  fine, 
shallow  punctures  on  vertex.  POL:OOL  = 
1:0.6.  Clypeus  1.2  X  as  wide  as  long. 
Mandible  simple,  apically  strongly  biden- 
tate. 

Thorax  microreticulate;  mesoscutum  and 
scutellum  with  only  scattered,  inconspicu- 
ous, very  shallow,  small  punctures;  prono- 
tum  and  mesopleuron  with  conspicuous, 
moderate-sized,  shallow  punctures.  Tegula 
1.5  X  as  long  as  wide,  the  posterior  inner 
corner  distinctly  inwardly  produced.  Pro- 
podeal  angles  evenly  rounded. 

Gaster  microreticulate  and  with  fine 
punctures. 

Male:  Coloration  as  in  female.  Parameres 
light  reddish-brown. 

Length  5.9-6.3  mm;  length  of  fore  wing 
3.6-4.3;  hamuli  7. 

Structurally  similar  to  female  but  differ- 
ing in  the  following  respects:  fore  femur 
(Fig.  5)  considerably  more  robust,  notched 
ventrally  in  basal  third  and  with  distally 
directed,  apically  rounded,  lamellate  pro- 
cess; tergum  VII  (Fig.  13)  with  surface 
flattened  medially,  with  hind  margin 
widely  rounded  and  medially  deeply  and 
narrowly  emarginate;  sternum  VII  with 
surface  convex  medially,  concave  laterally, 


with  apical  margin  widely  trilobed,  lateral 
lobes  ventrally  curved.  Genitalia  very  large 
(1.5  mm  long;  i.e.  half  the  length  of  the 
gaster);  outer  ramus  of  parameres  broad  in 
dorsal  view,  apically  obliquely  truncate  and 
densely  covered  with  fine,  long  setae;  inner 
ramus  proximally  of  varying  width  and 
distally  progressively  narrowing  and  mark- 
edly and  evenly  downcurved  to  form 
a  sharp,  well  sclerotized  hook  attaining  level 
of  lateral  posterior  angle  of  outer  ramus. 

Etymology. — The  name  femorata  serves  to 
draw  attention  to  the  uniquely  modified 
front  femur  of  the  male. 

Material  examined. — Holotype:  j,  NAMIBIA: 
1 1  km  S  of  Swakopmund  on  inland  side  of  road 
B2  to  Walvis  Bay  (22.46S  14.32E),  7.iv.2002  (F. 
W.  and  S.  K.  Gess)  [AMG].  Paratypes:  NAMI- 
BIA: same  data  as  holotype,  6  99,  13  $  $  [AMG]; 
same  data  as  holotype  but  date  14. iv. 2002,  1  9,  2 
$3  [AMG];  same  data  as  holotype  but  date 
20.iv.2002,  1  9,  4  S3  [AMG];  same  data  as 
holotype  but  date  30.iii.2004,  1  9,  2  J  3  [AMG]; 
same  data  as  holotype  but  date  31  .iii.2004 
[AMG],  1  9  [AMG];  Walvis  Bay,  22.ii.1990  (W. 
J.  Pulawski),  6  99,  2  3S)  [CAS].  (All  specimens 
collected  by  F.  W.  and  S.  K.  Gess  were  visiting 
the  pink  flowers  of  Trianthema  hereroensis  Schinz 
(Aizoaceae:  non-Mesembryanthema)  or  were  on 
the  sand  immediately  next  to  these  plants  where 
resting  or  mating.) 

Geographic  distribution. — Q.  femorata  is 
known  only  from  Namibia,  from  a  single 
locality  on  the  seaward  side  of  the  coastal 
dunes  at  the  northern  extremity  of  the 
Southern  Namib  of  Giess  (1971). 

Floral  associations. — Q.  femorata  has  con- 
sistently been  found  to  be  associated  solely 
with  Trianthema  hereroensis  Schinz  (Aizoa- 
ceae: non-Mesembryanthema). 

Nesting. — Unknown;  probably  in  the 
sand  beneath  the  hummock  forming  Tri- 
anthema bushes. 

Quartinia  geigeriae  Gess,  new  species 

Diagnosis. — Medium  sized  to  large  (3.8- 
5.0  mm).  Fore  wing  with  Cula  and  2m-cu 
complete  and  as  thick  as  other  veins. 
Tegula  short,  laterally  rounded,  with  pos- 


230 


Journal  of  Hymenoptera  Research 


terior  inner  corner  a  near  right  angle.  Both 
sexes  with  angles  of  propodeum  very 
markedly  posteriorly  produced,  lamellate 
and  subhyaline.  Female  with  head  and 
mesosoma  black,  tegulae  and  gaster  red- 
dish brown.  Male  with  head,  mesosoma 
and  gaster  black  with  yellowish-white 
markings. 

Description. — Female:  Black.  The  follow- 
ing are  various  shades  of  reddish  brown: 
labrum;  distal  two  thirds  of  mandibles; 
tegula;  scutellar  lamella;  median  section  of 
metanotum;  in  some  specimens  a  narrow 
streak  dorsally  on  outer  aspect  of  lamellate 
propodeal  angle  (rest  of  lamella  subhya- 
line); terga  I-IV  or  V  (narrow  posterior 
bands  lighter  in  colour  than  rest  of  terga). 
Underside  of  antenna,  distal  quarter  of 
femur,  entire  tibia  and  all  tarsomeres  of  all 
legs  light  reddish  yellow.  Wings  hyaline; 
veins  brown. 

Length  4.6-5.0  mm  (average  of  6:4.8 
mm);  length  of  fore  wing  2.7-3.0  (average 
of  6:2.9  mm);  hamuli  5;  length  of  extended 
tongue  3.1-3.2  mm. 

Head  in  front  view  1.23  X  as  wide  as 
long,  microreticulate  but  shiny,  with  sepa- 
rated, moderate  sized  punctures.  PO- 
L:OOL  -  1:0.85.  Clypeus  1.6  X  as  wide  as 
long  (length  measured  to  bottom  of  emar- 
gination;  1.36  X  if  measured  to  level  of 
antero-lateral  angles),  markedly  raised  an- 
teriorly and  laterally,  a  little  flattened 
medially;  anterior  margin  deeply  and 
evenly  emarginate;  antero-lateral  angles 
narrowly  rounded,  lamellate,  subhyaline. 

Mesosoma  microreticulate  but  shiny; 
mesonotum  and  scutellum  with  punctures 
slightly  larger  and  sparser  than  on  head; 
pronotum  with  punctures  similar  to  those 
on  head;  mesopleuron  with  punctures 
close  together,  reticulate-punctate  ventral- 
ly.  Propodeum  dorso-laterally  markedly 
raised,  dorso-medially  depressed  to  expose 
metanotum,  posteriorly  with  upper  three 
quarters  flat,  closely  reticulate-punctate 
and  lower  quarter  unpunctured  and  shiny, 
laterally  with  a  smooth,  shiny  depression 
and   arising  from   it  a   very  pronounced 


posteriorly  directed  lamella;  lamella  flat, 
very  thin,  subhyaline,  basally  slightly 
rugose  but  elsewhere  smooth,  marginally 
widely  and  evenly  rounded. 

Gaster  microreticulate  but  shiny;  punc- 
tures finer  and  shallower  than  on  head  and 
mesosoma,  becoming  progressively  finer 
posteriorly. 

Vestiture  generally  very  short  and 
sparse,  longer  and  more  noticeable  on 
labrum,  posterior  flat  surface  of  propo- 
deum and  declivous  anterior  face  of 
tergum  I. 

Male:  Black.  The  following  are  yellowish- 
white:  base  of  labrum  (in  some  specimens 
only);  clypeal  disk  and  adjoining  it  a  large 
medial  marking  on  frons  together  forming 
an  hour-glass-like  figure);  scape,  pedicel 
and  proximal  flagellomeres;  anterior  mar- 
gin of  pronotum  (transverse  band  in  some 
specimens  medially  interrupted  and  re- 
duced to  two  spots);  tegula  (except  for  pale 
testaceous  discal  spot);  in  some  specimens 
a  narrow  streak  dorsally  on  outer  aspect  of 
lamellate  propodeal  angle  (rest  of  lamella 
subhyaline);  narrow  posterior  bands  on 
terga  I-VI  (very  narrowly  anteriorly  wid- 
ened medially  on  II— VI;  immediate  vicinity 
of  emargination  of  tergum  VII;  distal  quar- 
ter of  femur,  entire  tibia  and  all  tarsomeres 
of  all  legs.  Varyingly  reddish  brown  are: 
mandible  distally;  concave  declivous  ante- 
rior surface  of  tergum  I.  Underside  of 
antennal  club  light  reddish,  upper  side 
brown.  Wings  hyaline;  veins  brown. 

Length  3.8-4.5  mm  (average  of  6:4.1 
mm);  length  of  front  wing  2.4-2.8  mm 
(average  of  5:2.6  mm);  hamuli  4-5. 

Structurally  very  similar  to  female  but 
puncturation  on  gaster  noticeably  coarser. 
Tergum  VII  reticulate  punctate,  postero- 
medially  with  a  shallow  V-shaped  emar- 
gination. Parameres  postero-laterally 
smoothly  curved  to  apex;  apex  not  hooked 
and  inner  edge  of  parameres  not  toothed. 
Labrum  shiny,  non-carinate.  Antenna  with 
poorly  defined,  elongate  club. 

Etymology. — The  name  geigeriae,  genitive 
singular,  is  formed  from  the  generic  name 


Volume  16,  Number  2,  2007 


231 


of  the  plants,  Geigeria  spp.  (Asteraceae),  on 
the  capitula  of  which  the  wasp  was  found 
foraging  for  nectar  or  nectar  and  pollen. 

Material  examined. — Holotype:  9,  NAMIBIA: 
Solitaire  (23.52S  16.00E),  30.iv.2002  F.  W.  and  S. 
K.  Gess)  (visiting  yellow  flowers  of  Geigeria 
ornativa  O.  Hoffrn.,  Asteraceae)  [AMG].  Para- 
types:  NAMIBIA:  same  data  as  holotype,  2  99,  6 
o'JlAMG];  between  Solitaire  and  Nomtsas 
(24.15S  16.33E),  l.v.2002  (F.  W.  and  S.  K.  Gess), 
10  99,  2  33  (9  99,  1  S  visiting  yellow  flowers  of 
Geigeria  ornativa;  1  9,  1  o  visiting  yellow  flowers 
of  Geigeria  pectidea  (DC.)  Harv.)  [AMG];  1  km  N 
of  Mariental  (24.37S  17.58E),  2.V.2002  (F.  W.  and 
S.  K.  Gess),  30  99,  4  33  (22  99,  2  33  visiting 
yellow  flowers  of  Geigeria  ornativa;  8  99,  2  33 
visiting  yellow  flowers  of  Geigeria  pectidea) 
[AMG];  between  Mariental  and  Keetmanshoop 
(24.54S  17.55E),  2.V.2002  (F.  W.  and  S.  K.  Gess),  1 
3  (visiting  yellow  flowers  of  Geigeria  pectidea) 
[AMG];  18  km  from  Ariamsvlei  on  road  to  Aroab 
[28.00S  19.43E],  14.V.1973  (C.  F.  Jacot-Guillar- 
mod),  5  99,  1  3  [AMG];  SOUTH  AFRICA: 
NORTHERN  CAPE:  Langvlei,  103  km  WNW  of 
Upington  [28.10S  20.16E],  14.V.1973  (C.  F.  Jacot- 
Guillarmod),  21  99,  2  33  [AMG]. 

Geographic  distribution. — Q.  geigeriae  is 
known  from  Namibia,  from  a  limited  area 
in  the  Semi-desert  and  Savanna  Transition 
(Escarpment  Zone)  and  the  adjoining 
Dwarf  Shrub  Savanna  of  Giess  (1971),  and 
from  a  closely  adjoining  locality  in  the 
Northern  Cape. 

Floral  associations. — Known  only  in  asso- 
ciation with  two  species  of  Geigeria,  Aster- 
aceae). 

Nesting. — Unknown. 

Discussion. — Q.  geigeriae  shares  with  Q. 
arteritis  Richards,  Q.  breyeri  Richards  and 
the  below  described  Q.  lameilata  the  pos- 
session of  markedly  backwardly  produced 
propodeal  lamellae.  Q.geigeriae  together 
with  breyeri  and  lameilata  is  readily  distin- 
guished from  artemis  in  having  the  poste- 
rior inner  corner  of  the  tegula  rounded  or 
a  near  right  angle,  not  markedly  produced 
inwards;  it  is  distinguished  from  both 
breyeri  and  lameilata  in  having  the  epicne- 
mium  rounded,  not  defined  by  a  low 
carina. 


Quartinia  lameilata  Gess,  new  species 

Diagnosis. — Large  to  very  large  (5.0- 
6.2  mm).  Fore  wing  with  Cula  and  2m-cu 
complete  and  as  thick  as  other  veins. 
Clypeus  raised  and  protruding  with,  espe- 
cially in  female,  marked  disto-lateral  lobes. 
Labrum  large,  very  noticeable,  in  female 
setose.  Epicnemium  defined  by  a  low 
carina.  Tegula  rounded  posteriorly,  with 
posterior  inner  corner  a  near  right  angle. 
Angles  of  propodeum  markedly  back- 
wardly produced,  lamellate. 

Description. — Female:  Black.  The  follow- 
ing are  yellowish-white:  in  some  speci- 
mens a  small  spot  on  disto-lateral  lobe  of 
clypeus;  transversely  oval  or  bilobed  me- 
dial marking  (in  some  specimens  reduced 
to  two  round  spots)  distally  on  frons 
immediately  above  clypeus;  in  a  single 
specimen  a  small  round  spot  in  ocular 
sinus;  broad  streak  behind  top  of  eye;  scape 
(distally),  pedicel,  intermediate  flagello- 
meres,  and  underside  of  antennal  club; 
pair  of  spots  on  dorsum  of  pronotum;  large 
mark  on  humeral  angle  (in  some  speci- 
mens remote  from  spots  on  dorsum,  in 
others  fused  with  them  to  form  a  continu- 
ous band);  variously  developed  streak  on 
postero-dorsal  angle  of  pronotum;  in  some 
specimens  a  small  spot  on  mesopleuron; 
tegula  (except  for  testaceous  median  area); 
in  some  specimens  a  small  streak  laterally 
(flanking  tegula)  on  mesonotum;  curved 
posterior  band  on  disk  of  scutellum;  angles 
of  propodeum;  posterior  bands,  reaching 
sides  and  generally  slightly  expanded 
medially  and  laterally,  on  terga  I-V;  apical 
half  of  tergum  VI;  postero-lateral  corners 
of  sterna  II— V  and  apical  half  or  more 
of  sternum  VI;  distal  half  or  less  of 
femur,  entire  tibia  and  tarsus  of  all 
legs.  Mandibles,  labrum  and  suffusion  on 
upper  surface  of  antennal  club  reddish- 
brown.  Wing  membrane  hyaline;  veins 
brown. 

Length  6.0-6.2  mm  (average  of  3:6.06 
mm);  length  of  fore  wing  3.9-4.08  mm 
(average  of  3:4.0  mm);  hamuli  4. 


232 


Journal  of  Hymenoptera  Research 


Head  in  front  view  1.21  X  as  wide  as 
long.  POL:OOL  =  1:0.83 

Clypeus  raised  and  protruding,  medially 
depressed,  distally  widely  and  deeply 
emarginate  and  with  marked  disto-lateral 
lobes.  Labrum  large,  longer  than  wide, 
apically  pointed,  setose.  Clypeus  and 
frons  moderately  shiny,  with  close, 
fairly  coarse  punctures  and  finely  micro- 
sculptured  interstices;  pronotum,  me- 
soscutum  and  scutellum  with  larger,  much 
more  sparsely  arranged  punctures  and 
extremely  finely  microsculptured  inter- 
stices; terga  uniformly  finely  punctured. 
Epicnemium  defined  by  a  low  carina. 
Tegula  rounded  posteriorly.  Angles  of 
propodeum  markedly  backwardly  pro- 
duced, at  mid-height  forming  a  rounded 
projection  and  below  that  translucently 
lamellate. 

Male:  Black.  The  following  are  yellow: 
clypeus  (other  than  for,  in  some  specimens 
including  holotype,  a  variously  sized  me- 
dian longitudinal  marking  and  in  all  speci- 
mens areas  immediately  adjacent  to  anten- 
nal  sockets);  large  transverse  marking 
distally  on  frons  immediately  above  clyp- 
eus; broad  streak  behind  top  of  eye;  scape 
(distally),  pedicel,  intermediate  flagello- 
meres  ,  and  underside  of  antennal  club; 
most  or  almost  entire  dorsal  surface  of 
pronotum  (except  in  all  specimens  small 
postero-lateral  area  flanking  tegula);  spot 
on  mesopleuron;  tegula  (except  for  testa- 
ceous median  area);  in  all  specimens 
a  marking  (ranging  from  a  minute  spot  to 
a  small  streak)  flanking  tegula  on  mesono- 
tum;  curved  posterior  band  on  disk  of 
scutellum;  scutellar  lamella;  angles  of 
propodeum;  posterior  bands  (anteriorly 
ill-defined  and  grading  into  reddish- 
brown),  reaching  sides  on  terga  I-VI  and, 
to  a  varying  degree,  apical  half  of  tergum 
VII;  ill-defined  posterior  bands  on  sterna 
II— VI;  most  of  sternum  VII;  distal  half  or 
less  of  femur,  entire  tibia  and  tarsus  of  all 
legs.  Mandibles  (wholly  or  in  part),  labrum 
and  suffusion  on  upper  surface  of  antennal 
club,  terga  and  sterna  anterior  to  posterior 


bands  reddish-brown.  Wing  membrane 
hyaline;  veins  brown. 

Length  5.0-5.8  (average  of  3:5.2  mm; 
holotype  5.0  mm);  length  of  fore  wing 
3.0  mm.  Head  in  front  view  1.24  X  as  wide 
as  long 

Structurally  very  similar  to  female  but 
puncturation  on  head  and  mesosoma 
markedly  coarser.  Tergum  VII  with  hind 
margin  shallowly  emarginate  and  postero- 
lateral lobes  rounded. 

Etymology. — The  name  lamellata  is  in- 
tended to  draw  attention  to  the  markedly 
backwardly  produced,  lamellate  angles  of 
the  propodeum. 

Material  examined.— Holotype:  3,  NAMIBIA, 
Rooibank  [23.11S  14.39E],  19.xii.1978  (H. 
Empey)  [AMG].  Paratypes:  NAMIBIA:  same 
data  as  holotype  but  date  28.xii.1978,  2  $$ 
[AMG];  Kaokoland  [Dist.],  Otjinungwa  (SE 
1712  Ab)  [17.17S  12.27E],  19-22.viii.1973  (? 
collector),  1  9  [NNIC];  Kaokoland  [Dist.], 
Khowarib  R.  (SE  1914  Ac)  [locality  not  pin- 
pointed], 17-19.V.1978  (S.  Louw,  M.-L.  Pen- 
rith), 1  ct  [NNIC];  Namib  Naukluft  Park, 
Vogelfederberg  (23.03S  15.00E),  21.  ii.  1988 
(G.  D.  Butler),  1  9  [NCP];  same  locality, 
24.L1988  (R.  Miller  and  L.  Stange),  1  9  [FSCA]; 
Luderitz  [Dist.],  Sossusvlei  (SE  2415  Cd)  [24.43S 
15.20E],  12-19.ix.1971  (?  collector),  3  99  [NNIC]; 
Luderitz  [Dist.],  Kanaan  104  (SE  2516  Cc) 
[25.50S  16.09E],  6-7.X.1972  (?  collector),  10  99 
[NNIC]; 

Geographic  distribution. — Quartinia  lamel- 
lata is  widespread  in  the  western  parts  of 
Namibia,  collection  localities  spanning 
eight  degrees  of  latitude  and  falling  in  the 
Mopane  Savanna,  Central  Namib  and 
Southern  Namib /Semi-desert  and  Savanna 
Transition  (Escarpment  Zone)  of  Giess 
(1971). 

Floral  associations. — Unknown. 

Nesting. — Unknown. 

Discussion. — See  discussion  under  geiger- 
iae.  On  the  basis  of  the  characters  there 
listed,  lamellata  is  closest  to  breyeri  but  may 
readily  be  distinguished  from  that  species 
by  its  larger  size,  differently  developed 
clypeus  and  labrum,  differences  in  punc- 
turation and  in  colour  pattern. 


Volume  16,  Number  2,  2007 


233 


ACKNOWLEDGMENTS 

The  following  individuals  are  thanked  for  much 
appreciated  assistance  as  specified:  Dr  Sarah  Gess  of 
the  Albany  Museum,  Grahamstown,  co-collector  of 
most  of  the  Albany  Museum's  Quartinia  material,  for 
over  thirty  years  of  happy,  productive  and  synergistic 
fieldwork,  for  valuable  discussion  and  encourage- 
ment; Mr  Robert  W  Gess  for  field  assistance  in 
southern  Namibia  and  the  Northern  Cape  in  1996; 
Mr  David  W  Gess,  Ms  Gaby  T  Gess  and  Miss  Gaby 
Maria  Gess  for  field  assistance  at  Melkbosstrand  in 
2005  and  at  Yzerfontein  in  2006;  Coleen  Mannheimer 
of  the  National  Herbarium  of  Namibia,  Windhoek  for 
her  invitation  to  join  the  Herbarium  party  on  their 
expeditions  to  the  Sperrgebiet  in  2002,  2003  and  2005 
and  also  for  her  determination  of  voucher  specimens 
of  Namibian  plants  visited  for  pollen  and  nectar  by 
masarines;  Eugene  Marais  of  the  Namibian  National 
Insect  Collection,  Windhoek,  Connal  Eardley  of  the 
National  Collection  of  Insects,  Pretoria,  Wojciech 
Pulawski  of  the  California  Academy  of  Sciences,  San 
Francisco,  and  Lionel  Stange  and  Jim  Wiley  of  the 
Florida  State  Collection  of  Arthropods,  Gainesville  for 
the  loan  of  specimens  from  their  respective  collec- 
tions; Caroline  Mayer  of  BIOTA-Southern  Africa, 
Hamburg  University  for  the  gift  of  specimens  collect- 
ed by  herself  in  Namaqualand. 

Grateful  thanks  are  expressed  to  all  those  bodies 
which  issued  permits  for  the  collection  of  insects  and 
plant  samples,  namely:  the  Namibian  Ministry  of 
Environment  and  Tourism;  the  Namibian  Ministry  of 
Mines  and  Energy  as  also  NAMDEB  (Pty)  Ltd  (for  the 
Sperrgebiet  -  Diamond  Area  No  1);  the  Department  of 
Nature  and  Environmental  Conservation,  Northern 
Cape;  CapeNature  (Western  Cape  Nature  Conserva- 
tion Board);  Department  of  Economic  Affairs,  Envi- 
ronment and  Tourism,  Eastern  Cape  (Western  Re- 
gion); and  the  Nature  Conservation  Division,  City  of 
Cape  Town  (for  the  Blaauwberg  Conservation  Area). 

Debi  Brody  of  the  Graphics  Services  Unit  of  Rhodes 
University,  Grahamstown  is  thanked  for  help  with  the 
production  of  the  figures. 

The  National  Research  Foundation  (NRF)  is 
thanked  for  running  expenses  grants  for  fieldwork 


during  the  course  of  which  much  of  the  present 
material  was  collected.  The  Board  of  Trustees  of  the 
Albany  Museum  is  thanked  for  Research  Contracts 
granted  to  the  author  and  Dr  Sarah  Gess  since  2003, 
which  have  given  them  continued  use  of  the  mu- 
seum's facilities  since  their  retirements. 

LITERATURE  CITED 

Andre,  Ed.  1884  Species  des  Hymenopteres  d'Europe  et 

Algerie.  Vol.  2.  Beaune,  Andre  and  Andre. 

Carpenter,  J.  M.  2001.  Checklist  of  species  of  the 
subfamily  Masarinae  (Hymenoptera:  Vespidae). 
American  Museum  Novitates  3325:  1-39. 

Gess,  F.  W.  and  S.  K.  Gess.  1999.  The  use  by  wasps, 
bees  and  spiders  of  shells  of  Trigonephrus  Pilsb. 
(Mollusca:  Gasteropoda:  Dorcasiidae)  in  desertic 
winter-rainfall  areas  in  southern  Africa,  journal  of 
Arid  Environments  43:  143-153. 

Giess,  W.  1971.  A  preliminary  vegetation  map  of 
South  West  Africa.  Dinteria  4:  1-114. 

Greathead,  D.  J.  1999.  Apolysis  sp.  (Diptera:  Bombylii- 
dae)  reared  from  Quartinia  sp.  (Hymenoptera: 
Vespidae:  Masarinae).  journal  of  Arid  Environ- 
ments 43:  155-157. 

.  2006.  New  records  of  Namibian  Bombyliidae 

(Diptera),  with  notes  on  some  genera  and 
descriptions  of  new  species.  Zootaxa  1149:  1-88. 

Leistner,  O.  A.  and  J.  W.  Morris.  1976.  Southern 
African  Place  Names.  Annals  of  the  Cape  Provincial 
Museums  (Natural  History)  12:  i-iv,  1-565. 

Richards,  O.  W.  1962.  A  revisional  study  of  the  masarid 
wasps  (Hymenoptera,  Masaridae).  London:  British 
Museum  (Natural  History). 

.  1982.  A  new  species  of  Quartinioides  Richards 

(Hymenoptera,  Masaridae.  Bollettino  del  Museo 
civica  di  storia  naturale  di  Venezia  32:  199-200. 

Schulthess,  A.  von.  1929.  Contribution  to  the 
knowledge  of  African  Masaridae  (Vespoidea). 
Annals  and  Magazine  of  Natural  History  (10)3: 
498-511. 

Vecht,  J.  van  der.  and  J.  M.  Carpenter.  1990.  A 
catalogue  of  the  genera  of  the  Vespidae  (Hyme- 
noptera). Zoologische  Verhandelingen  260:  1-62. 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  234-265 

Torymidae  (Hymenoptera:  Chalcidoidea)  Associated  with  Bees 
(Apoidea),  with  a  List  of  Chalcidoid  Bee  Parasitoids 

E.  E.  Grissell 

Systematic  Entomology  Laboratory,  PSI,  Agricultural  Research  Service, 
U.  S.  Department  of  Agriculture,  c/o  National  Museum  of  Natural  History,  Smithsonian  Institution, 

P.O.  Box  37012,  MRC  168,  Washington,  D.  C.  20013-7012 


Abstract. — Thirty-one  species  of  Torymidae  (Hymenoptera:  Chalcidoidea)  are  associated  with 
bees.  In  this  review  each  is  keyed  and  discussed,  and  geographic  ranges  and  hosts  are  given.  Most 
species  are  illustrated.  Torymids  represent  about  one-fourth  of  the  135  species  of  Chalcidoidea 
associated  with  bees.  Two  summary  lists  are  presented  for  all  chalcidoids,  including  Torymidae, 
and  the  216  bee  species  with  which  they  are  associated.  One  is  arranged  as  a  bee/parasitoid  list  and 
the  other  as  a  parasitoid/bee  list. 


Considering  that  22,000  species  of  Chal- 
cidoidea (Noyes  2003)  and  16,000-17,000 
bee  species  (Michener  2000)  have  been 
described,  the  number  of  chalcidoids  re- 
ported associated  with  bees  is  surprisingly 
small.  At  most  135  different  chalcidoids 
have  been  reared  from,  or  associated 
with,  216  bee  species  (see  Appendix,  de- 
rived from  Noyes  2003).  Of  these,  the 
families  Torymidae  and  Leucospidae 
have  the  highest  percentage  of  the  bee 
parasitoids  (each  22-23%),  followed 
closely  by  Pteromalidae  (18%).  The  other 
families  associated  with  bees  are:  En- 
cyrtidae  (13%),  Eulophidae  (13%),  Chalci- 
didae  (5%),  Eurytomidae  (5%),  Eupelmi- 
dae  (3%),  Mymaridae  (0.6%),  and  Perilam- 
pidae  (0.6%)  (Appendix:  based  on  Noyes 
2003). 

Although  Torymidae  and  Leucospidae 
have  the  highest  number  of  bee  parasitoids 
among  Chalcidoidea,  this  figure  is  some- 
what misleading.  Of  approximately  1,000 
torymid  species,  only  31  are  known  (or 
suspected)  to  attack  bees  (Grissell  1995, 
2000,  2005;  Noyes  2003),  so  a  predilection 
for  bee  hosts  is  not  especially  pronounced 


in  the  family.  The  host  range  of  this  family 
is  extremely  broad,  but  nearly  80%  of  the 
known  hosts  are  shared  equally  between 
the  Hymenoptera  and  Diptera,  most  of 
which  are  gall-forming  cynipids  and  ceci- 
domyiids  (Grissell  1995).  Conversely,  the 
entire  family  Leucospidae,  consisting  of 
135  species,  has  been  presumed  to  parasit- 
ize aculeate  Hymenoptera — solitary  bees, 
and  less  frequently,  solitary  wasps.  In 
reality,  however,  hosts  are  known  only 
for  about  30  leucospid  species  (Boucek 
1974,  Noyes  2003),  so  the  true  relationship 
of  the  family  to  bees  is  largely  unknown. 
Recently  a  species  of  leucospid  was  re- 
ported as  an  ectoparasitoid  of  an  ichneu- 
monid  attacking  a  cerambycid  in  limbs  of 
apricot  in  Iran  (Hesami  et  al.  2005).  This 
finding  casts  doubt  on  our  concept  of  host 
specificity  in  Leucospidae. 

In  this  paper  I  present  a  summary  of 
torymid  species  reported  to  attack  bees, 
including  a  review  of  published  informa- 
tion for  each  species  and  a  key.  I  also 
include  a  world  bee/chalcidoid  and  chal- 
cidoid/bee  list  for  all  Chalcidoidea  re- 
portedly associated  with  bees  (Appendix). 


Volume  16,  Number  2,  2007 


235 


As  with  many  chalcidoid  records,  the  true 
host  is  not  always  indicated  by  the  host 
record  given  (Noyes  1994).  Many  bee  host 
records  are  simply  nest  rearings  and  may 
have  been  contaminated  by  other  true 
parasitoids,  cleptoparasitoids,  inquilines, 
and  simple  space  usurpers  of  all  sorts, 
many  not  even  hymenopteran.  Similarly, 
a  mud  wasp's  nest  may  be  usurped  by 
a  nesting  bee,  thus  causing  confusion  as  to 
the  true  host  (Rust  1974).  Bee  nests,  as  well 
as  almost  any  other  ecological  niche,  offer 
complex  arrays  of  hosts,  many  of  which  are 
not  even  suspected  at  the  time  of  rearing. 
For  example,  Glypkomerus  stigma  (Fabri- 
cius)  was  reported  from  Melitoma  taurea 
(Say)  (Apidae),  but  this  is  likely  to  be  an 
error  because  all  other  records  for  species 
of  Glyphomerus  are  gall-forming  cynipids  or 
rarely  eurytomids  (Grissell  1995).  With 
respect  to  bee  parasitoids,  therefore,  all 
records  should  be  considered  tentative 
until  established  by  dissection  and  obser- 
vation. Within  the  Torymidae  listed  in  this 
paper,  I  point  out  that  several  are  likely  not 
to  be  true  bee  parasitoids.  In  those  few 
cases  where  the  biologies  of  torymids  are 
known  they  are  generally  solitary,  idiobio- 
tic  larval  ectoparasitoids,  but  in  several 
genera  (e.g.,  Monodontomerus,  Microdonto- 
merus)  larvae  are  known  to  be  gregarious 
(Grissell  2000,  2005). 

In  examining  host  records  presented  in 
the  Appendix  several  reviewers  suggested 
that  it  might  be  informative  to  summarize 
parasitoid  data  with  respect  to  bee  biology 
as  there  appeared  to  be  a  bias  towards  twig 
and  cavity  nesting  bees,  with  ground- 
nesters  being  under-represented.  I  solicited 
the  input  of  two  recognized  bee  authori- 
ties: Frank  Parker,  who  specializes  in  twig- 
nesters,  and  Jerry  Rozen,  who  specializes 
in  ground-nesters,  and  both  agreed  that  the 
data  suggested  cavity  nesters  were  the 
predominant  host  representatives.  These 
are  primarily  twig  nesters,  bees  that  nest  in 
pre-existing  crevices  or  cavities,  and  bees 
that  re-use  old  bee  nests.  Some  of  these 
nests  may  be  external,  for  example  resin 


nests  attached  to  objects  such  as  twigs  and 
rocks.  According  to  Rozen  most  of  the 
records  are  indicative  of  shallow  nesting 
bees,  and  he  suggested  that  ground  nesting 
bees  in  general  would  be  less  likely  to 
harbour  parasitoids  because  they  might 
have  a  more  difficult  time  entering  nests 
and  crawling  down  the  "...  long,  main 
tunnels"  to  find  their  host.  He  also  pointed 
out  that  old  bee  nests  and  shallow  cavities 
are  frequently  re-used  several  times,  thus 
encouraging  the  build-up  of  large  parasit- 
oid populations.  Parker  suggested  that 
twig-nesting  bees  are  more  likely  to  be 
sampled  because  they  readily  come  to 
artificial  traps  set  out  by  the  collector.  They 
are  also  easier  to  extricate  and  study  in 
these  nests.  Conversely,  ground  nesting 
bees  must  be  actively  hunted  by  the 
collector,  are  less  easily  found,  and  require 
painstaking  excavation  to  reveal  nest  de- 
tails. 

In  general,  then,  records  summarized  in 
the  Appendix  indicate  that  host  data  are 
biased  towards  parasitoids  attacking  cavity 
nesting  bees  and  that  multiple  causes 
contribute  to  this  bias.  Whatever  cursory 
glimpses  the  bee/parasitoid  host  list  may 
reveal,  and  considering  the  numerical  size 
of  the  chalcidoid  and  apoid  groups,  it 
appears  that  much  remains  to  be  discov- 
ered. Within  existing  literature,  relatively 
little  is  devoted  to  parasitization  and  then 
primarily  only  to  a  few  solitary  bee  species 
(e.g.,  the  alfalfa  leafcutting  bee,  Stolbov  et 
al.  1986),  whereas  with  few  exceptions 
(e.g.,  Zerova  and  Romasenko  1986)  there 
is  scarcely  any  comprehensive  literature 
pertaining  to  solitary  bee  parasitoids. 

METHODS 

In  the  following  discussions  host  names 
are  given  without  authors.  Complete 
authors'  names  may  be  found  in  the 
Appendix.  Within  discussions,  hosts  are 
listed  alphabetically  by  family,  but  in  the 
host  listing  all  hosts  are  alphabetic  regard- 
less of  family. 


236  Journal  of  Hymenoptera  Research 


KEY  TO  TORYMIDAE  ASSOCIATED  WITH  SOLITARY  BEES 

1  Anterior   edge   of   metapleuron    straight,    not   projecting    forward    as    lobe    into 

mesepimeron  (Fig.  2) 4 

—  Anterior  edge  of  metapleuron  (usually  its  upper  half)  projecting  forward  as  lobe  into 
mesepimeron  (Fig.  1),  which  is  subdivided  into  upper  and  lower  sections,  lower 
section  delimited  by  anterior  groove   Torymus  Dalman  2 

2  Hind  coxa  dorsally  covered  with  short  setae  (Fig.  3),  coarsely  reticulate;  propodeum 

areolate-rugose,   heavily  carina te   (Fig.  5);   frenal   area   less   than   1/5   length   of 

scutellum  (Fig.  7) 3 

Hind  coxa  dorsally  bare  (a  few  long  setae  may  be  present;  Fig.  4),  smooth  and 
polished;  propodeum  essentially  smooth  (Fig.  6);  frenal  area  1/3  to  almost  1/2 

length  of  scutellum  (Fig.  8)  (Palearctic,  Australasian  [?introduced]) 

Torymus  armatus  Boheman 

3(2)         Head  dorsum,  mesosoma,  and  hind  coxa  coppery  with  greenish  tints;  at  least  part  of 

hind  femur  orange,  concolorus  with  tibia  (Palearctic)    .  .  .      Torymus  cupreus  (Spinola) 
Head  dorsum,  mesosoma,  and  hind  coxa  metallic  green  or  blue;  entire  hind  femur 

metallic  green  or  blue,  contrasting  with  orange  tibia  (Nearctic) 

Torymus  zabriskii  (Cresson) 

4(1)  Fore  wing  with  marginal  and  stigmal  veins  conspicuously  thickened  relative  to 
submarginal  vein,  postmarginal  vein  not  projecting  beyond  tip  of  stigmal  vein 
(Figs  9,  12,  13),  and  with  marginal  vein  slightly  removed  from  margin  of  wing 
(Fig.  13;  may  be  somewhat  difficult  to  see);  malar  distance  longer  than  intermalar 
distance    (Figs  14,    15);    mandibles    reduced,   scarcely   visible,    tips   not   meeting 

medially  when  closed,  apically  without  teeth    Echthrodape  Burks  5 

Fore  wing  with  marginal  and  stigmal  veins  not  conspicuously  thickened  relative  to 
submarginal  vein,  with  postmarginal  vein  longer  than  stigmal  vein  (Fig.  10),  and 
with  marginal  vein  at  edge  of  wing  margin;  malar  distance  subequal  to  or  shorter 
than  intermalar  distance  (Fig.  11);  mandibles  visible,  tips  meeting  medially  when 
closed,  apically  with  teeth   6 

5(4)  Postmarginal  vein  developed,  longer  than  stigmal  vein,  which  is  slender  and  petiolate 
(Fig.  12);  genae  straight,  not  concave  (Fig.  14)  [Papua  New  Guinea,  Austra- 
lia]       Echthrodape  papuana  Boucek 

Postmarginal  vein  reduced,  subequal  to  stigmal  vein,  which  is  thick  and  sessile 
(Fig.  13);  genae  concave  (Fig.  15)  [Kenya]    Echthrodape  africana  Burks 

6(4)  Occipital  carina  absent  (Fig.  16),  weakly  or  questionably  developed,  or  if  apparent, 
then  medially  arched  and  midway  between  hind  ocelli  and  occipital  foramen  and 
not  reaching  hypostomal  carina  (Fig.  17)  (head  usually  vertical  with  dorsoposterior 
aspect  slightly  concave  and  the  carina,  if  present,  easily  seen);  hind  femur  slender, 
apicoventrally  either  without  tooth  (Fig.  34),  angulate,  or  vaguely  serrate; 
metasomal    terga    with   or   without   apicomedian   emarginations,    often    weakly 

sclerotized 26 

Occipital  carina  well  developed,  dorsal  margin  not  greatly  arched  but  nearly 
horizontal  (Fig.  18),  closer  to  occipital  foramen  than  to  hind  ocelli  and  reaching 
hypostomal  carina  (head  usually  tilted  forward  with  dorsoposterior  aspect 
conspicuously  concave  and  occipital  carina  easily  seen,  but  head  must  be  removed 
to  see  hypostomal  carina);  hind  femur  apicoventrally  with  abrupt  tooth  (Figs  35,  37, 
38),  or  greatly  swollen  and  angulate  (Fig.  36);  metasomal  terga  heavily  sclerotized, 

without  apicomedian  emarginations  [Holarctic,  Neotropical,  Oriental]    

Monodontomerus  Westwood  7 


Volume  16,  Number  2,  2007  237 

7(6)  First  2  flagellar  segments  reduced  in  length,  ring-like  (Fig.  19);  hind  femur  swollen 
with    distal    subapical    angle    but    without    distinct    tooth    (Fig.  36)    [Nearc- 

tic] Monodontomerus  thorpi  Grissell 

At  most,  first  flagellar  segment  reduced  in  length  (Fig.  20);  hind  femur  relatively 
narrow  with  distinct  subapical  tooth  (Figs  37,  38)    8 

8(7)  Female,  face  transverse,  intermalar  distance  3.5  to  5x  length  of  malar  distance;  male, 
face  grotesquely  modified,  entirely  sunken  medially  (as  if  entirely  consisting  of 
scrobal  basin)  (Fig.  22),  with  sharp  edge  mesad  of  eye  (Fig.  21)  [Palearctic,  Nearctic 

(introduced)]    Monodontomerus  osmiae  Kamijo 

Both  sexes,  face  at  most  slightly  transverse,  intermalar  distance  from  1  to  3X  length  of 
malar  distance;  male  with  face  not  medially  sunken,  scrobal  basin  normal,  though 
areas  on  either  side  of  scrobe  may  be  slightly  depressed 9 

9(8)         Clypeus  greatly  elongate  (Figs  23,  24)  [Palearctic]     .  .    Monodontomerus  anthidiorum  Lucas 
Clypeus  either  barely  reaching  to  or  beyond  line  drawn  across  lateral  corners  of  oral 
fossa  (Figs  25-28)     10 

10(9)       Upper  mesepimeral  area  with  anterior  half  reticulately  sculptured  and  anterodorsal 

corner  diagonally  striate  extending  nearly  to  transepimeral  sulcus  (Fig.  53)     ....      11 
Upper  mesepimeral  area  nearly  entirely  polished  with  striae  scarcely  extending  half 
way  to  transepimeral  sulcus  (Figs  54,  55)     12 

11(10)  Discal  setae  of  fore  wing  not  extending  into  basal  area  (as  in  Fig.  31);  female  with 
ovipositor  sheaths  shorter  than  metasoma;  male  with  clypeus  recessed  (not 
extending  beyond  line  drawn  across  lateral  corners  of  oral  fossa),  malar  sulcus 
absent  or  obscure,  malar  distance  subequal  to  intermalar  distance;  scape  with 
ventral   surface   slightly   keeled    vertically    (i.e.,   not   flat),   no   pores   visible 

[Palearctic]     Monodontomerus  laticornis  Grissell  and  Zerova 

Discal  setae  of  fore  wing  extending  into  basal  area  (as  in  Fig.  29);  female  with  ovipositor 
sheaths  as  long  as  or  longer  than  entire  body;  male  with  clypeus  extending  beyond 
line  drawn  across  lateral  corners  of  oral  fossa,  malar  sulcus  present;  malar  distance 
about  1.5X  intermalar  distance;  scape  with  ventral  surface  flat,  covered  with  pores 
visible  at  100 X  [Nearctic] Monodontomerus  dementi  Grissell 

12(10)     Metasomal  tergum  2  dorsally  with  reticulate  to  strigate  sculpture  in  distal  half     ...      13 

—  Metasomal  tergum  2  dorsally  smooth,  polished  in  distal  half     14 

13(12)  Distal  portion  of  postmarginal  vein  equal  in  length  to  proximal  portion  (as  in  Fig.  30); 
rim  of  scutellum  apically  widened,  somewhat  projecting;  females,  metasomal 
tergum  6  acute  in  profile  (as  in  Fig.  32);  male,  fore  leg  unmodified  (i.e.,  normal)  (as 
in  Fig.  39),  tibia  equal  in  length  to  femur  and  not  ventrobasally  concave,  tarsomeres 

elongate  (claw  length  equal  to  or  shorter  than  tarsomere  4)  [Nearctic]     

Monodontomerus  dianthidii  Gahan 

Distal  portion  of  postmarginal  vein  0.33 X  length  of  proximal  portion  (as  in  Fig.  31); 
rim  of  scutellum  apically  even  in  width,  not  projecting;  female,  metasomal  tergum 
6obtuse  in  profile  (as  in  Fig.  33);  male,  fore  leg  modified  (Fig.  40),  tibia  shorter  in 
length  than  femur  and  ventrobasally  concave,  tarsomeres  shortened  (claw  length 
equal  to  tarsomeres  3  and  4)  [Nearctic]    Monodontomerus  breincrus  Grissell 

14(12)  Malar  sulcus  absent  (Fig.  28),  or  if  weakly  apparent,  greatly  curved  backward  from 
lower  margin  of  eye  then  curving  downward   to  join  edge  of  malar  opening 

(Fig.  27);  lower  face  protuberant  in  profile  (Fig.  28)  [Nearctic] 

Monodontomerus  bakeri  Gahan 

Malar  sulcus  well  developed,  straight  (Figs  25-26),  or  slightly  curved  from  lower 
margin  of  eye  to  lateral  edge  of  malar  opening;  lower  face  flat  (not  bulging)  in 
profile  (Fig.  26)    15 

15(14)  Frenal  area  medially  highly  polished,  appearing  glabrous,  faint  coriaceous  sculpture 
may  be  seen  with  difficulty  at  some  angles  of  view  (questionable  species  will  run 
through  either  couplet  of  key) 16 


238  Journal  of  Hymenoptera  Research 

Frenal  area  medially  sculptured,  may  be  uniformly  similar  overall  or  relatively  less 
prominent  than  laterally,  never  glabrous,  sculpture  easily  visible  at  any  angle  of 

view  (questionable  species  will  run  through  either  couplet)     19 

16(15)  Costal  cell  above  with  apical  setal  row  incomplete,  confined  to  distal  1  /2  or  less  of  cell 
(as  in  Fig.  31);  female,  metasomal  tergum  6  strongly  concave  in  profile  (as  in 
Fig.  32)  [Nearctic]    Monodontomerus  torchioi  Grissell  (most  specimens) 

—  Costal  cell  above  with  apical  setal  row  complete  (as  in  Fig.  29);  female,  metasomal 

tergum  6  weakly  concave  in  profile  (as  in  Fig.  33) 17 

17(16)  Frenal  area  apicomedially  intruding  into  rim,  punctures  of  rim  reduced  in  size  at  point 
of  intrusion  (Fig.  42);  stigma  and  uncus  relatively  short,  postmarginal  vein  with 

proximal  and  distal  section  subequal  in  length  (Fig.  30)  [Holarctic]     

Monodontomerus  aeneus  (Fonscolombe) 

—  Frenal  area  with  apical  rim  not  interrupted  posteriorly  at  median  margin,  punctures  of 

rim  as  large  or  larger  at  apex  as  on  sides  (as  in  Fig.  41);  stigma  and  uncus  elongated, 
postmarginal  vein  with  proximal  section  longer  than  distal  (as  in  Fig.  31)     18 

18(17)  Female  ovipositor  sheaths  swelling  distally  (i.e.,  not  parallel-sided);  male  hind  femur 
broad,  widening  apically,  about  2.5  X  as  long  as  wide  (Fig.  38)  [Palearctic]   .... 

Monodontomerus  rugulosus  Thomson 

Female  ovipositor  sheaths  same  width  throughout  (i.e.,  parallel-sided);  male  hind 
femur  narrow,  dorsal  and  ventral  margins  essentially  parallel  (Fig.  37),  about  3.5  X 
as  long  as  wide  [Neotropical]    Monodontomerus  argentinus  Brethes 

19(15)     Costal  cell  above  with  apical  setal  row  complete  (Fig.  29)    20 

Costal  cell  above  with  apical  setal  row  incomplete,  confined  to  distal  1/3  to  1/2  of  cell 
(Fig.  31)  or  appearing  absent  (2  or  3  setae  may  be  present  at  apex  as  in  Figs  43,  44)   ...     22 

20(19)     Scape  about  4x  longer  than  wide,  greater  in  length  (about  1.3X)  than  distance  from 

venter  of  torulus  to  apical  margin  of  clypeus  [Nearctic,  Neotropical]     

Monodontomerus  mexicanus  Gahan 

Scape  about  3X  longer  than  wide,  subequal  in  length  to  distance  from  venter  of 
torulus  to  apical  margin  of  clypeus 21 

21(19)  Stigma  rectangular,  proximally  elongated  towards  base  of  wing  (Figs  51,  52); 
postmarginal  vein  with  distal  length  less  than  proximal  length  (Figs  51,  52);  male 

face  with  depression  laterad  of  scrobal  basin  [Nearctic]     

Monodontomerus  acrostigmus  Grissell 

Stigma  squarish,  neither  stigma  nor  proximal  angle  elongated  (as  in  Figs  10,  44); 
postmarginal  vein  with  distal  length  subequal  to  basal  length  (as  in  Fig.  44);  male  face 
convex  laterad  of  scrobal  basin  [Holarctic]    Monodontomerus  obscurus  Westwood 

22(19)  Admarginal  setae  reaching  bases  of  marginal  vein  and  parastigma  (Fig.  43);  intermalar 
distance  subequal  to  3x  malar  distance  (Fig.  48);  both  mandibles  with  single  apical 

tooth  and  small  secondary  tooth  on  dorsal  margin  (Fig.  48)  [Nearctic]    

Monodontomerus  mandibularis  Gahan 

Admarginal  setae  either  not  reaching  base  of  marginal  vein  or  apex  of  parastigma 
(Fig.  44);  intermalar  distance  less  than  2.5 X  malar  distance  (Fig.  47);  both  mandibles 
with  2  apical  teeth,  and  small  third  tooth  on  dorsal  margin  (Figs  47)    23 

23(22)  Transepimeral  sulcus  incomplete  (Figs  53,  55);  upper  anterior  margin  of  costal  cell 
with  setal  row  in  apical  1/4  to  1/3  (as  in  Fig.  31);  male,  scape  in  side  view  slightly 
curved  in  profile  (Fig.  56),  area  beneath  torulus  flat,  sculptured,  and  setose  ....  24 
Transepimeral  sulcus  complete  (Fig.  54),  appearing  as  a  sculptured  groove;  upper 
anterior  margin  of  costal  cell  with  1  to  3  setae  at  apex  (Figs  43,  44);  male,  scape  in 
lateral  view  strongly  C-shaped  (Figs  57,  58),  area  beneath  torulus  slightly  swollen, 
polished,  and  asetose     25 

24(23)  Frenal  area  medially  with  reticulate  sculpture  readily  apparent,  area  may  be  shiny,  but 
sculpture  visible  at  any  angle  of  view;  male,  ventral  surface  of  scape  without  pores 
visible  at  100X  [Nearctic]     Monodontomerus  montivagus  Ashmead 


Volume  16,  Number  2,  2007  239 

Frenal  area  medially  with  reticulate  sculpture  visible  only  at  some  angles  of  view  and 
seen  only  with  difficulty,  area  shiny  and  appearing  polished;  male,  ventral  surface 
of  scape  evenly  covered  with  pores  easily  visible  at  100X  [a  few  atypical  specimens 
run  here,  but  most  to  couplet  16  based  on  the  polished  frenal  area] 
[Nearctic]     Monodontomerus  torchioi  Grissell 

25(23)  Females,  ovipositor  subequal  to  metasoma  (ca.  1-1. 2X);  scape  orange  to  yellow 
without  metallic  infusion  especially  ventrally;  male,  scape  greatly  laterally 
compressed  (ventral  and  dorsal  surfaces  essentially  absent),  outer  surface  flat, 
polished,  asetose,  and  curving  smoothly  to  inner  surface  without  interruption 

(Fig.  57),  no  pores  visible  at  100x  [Nearctic]    Monodontomerus  parkeri  Grissell 

Females,  ovipositor  obviously  longer  than  metasoma  (ca.  1.5-1.8X);  scape  with 
metallic  green  infusion  at  least  ventrally  (may  be  complete  or  confined  to  area  just 
beneath  pedicel);  male,  scape  dorsoventrally  compressed,  curved,  with  dorsal  and 
ventral  surfaces  parallel  and  delimited  by  right-angled  edge  (Fig.  58),  ventral 
surface  polished  and  covered  with  pores  visible  at  100  x  though  difficult  to  see 
[Nearctic] Monodontomerus  tepedinoi  Grissell 

26(6)  Marginal  vein  long,  3-7X  length  of  postmarginal  vein  and  at  least  6X  length  of  stigmal 
vein;  occipital  carina  present,  its  lateral  edges  extending  at  least  in  line  with  dorsum 

of  hypostomal  foramen  [Oriental]    Pseudotorymus  indicus  (Mani) 

Marginal  vein  short,  1-2.5  X  length  of  postmarginal  vein  and  2-5  X  length  of  stigmal 
vein  (Figs  45,  46);  occipital  carina  absent  (Fig.  16)  or,  if  present,  its  lateral  edges  not 
(or  scarcely)  extending  in  line  with  venter  of  occipital  foramen  (Fig.  17)    27 

27(26)     Occipital  carina  absent  (Fig.  16)    Microdontomerus  Crawford  28 

Occipital  carina  visible  in  dorsal  view  as  finely  polished  line  raised  distinctly  above 

surface  sculpture  (as  in  Fig.  17)  [Palearctic] 

Adontomerus  Nikol'skaya  (A.  gregalis  (Steffan)  and  A.  nesterovi  Zerova) 

28(27)  Fore  wing  setation  (Figs  45,  46)  reduced;  basal  cell  open  behind,  i.e.,  cubital  vein 
basally  at  most  with  few  isolated  setae;  basal  vein  at  most  with  isolated  setae;  basal 

cell  without  distinct  setal  row  paralleling  submarginal  vein    29 

Fore  wing  setation  (as  in  Figs  29,  31)  not  reduced  (except  in  admarginal  area  of  some 
species):  basal  cell  closed  behind,  i.e.,  cubital  vein  essentially  completely  setose  to 
base  of  wing;  basal  vein  with  distinct  setal  row  and  basal  cell  with  setal  row 
paralleling  nearly  entire  submarginal  vein     30 

29(28)  Postmarginal  vein  (Fig.  46)  about  0.75  X  as  long  as  marginal  vein;  fore  wing  with 
admarginal    area    (Fig.  46)    not   well    defined    posteriorly   by   setal    line,    with 

admarginal  setae  nearly  as  uniform  as  central  area  of  wing  [Nearctic]    

Microdontomerus  enigma  Grissell 

Postmarginal  vein  (Fig.  45)  about  0.5 X  as  long  as  marginal  vein;  fore  wing  with 
admarginal  area  (Fig.  45)  well-defined  posteriorly  by  setal  line,  with  few  sparse  setae 
not  as  uniform  as  central  area  of  wing  [Nearctic]   ....     Microdontomerus  parkeri  Grissell 

30(28)     Eye  height  nearly  3x  malar  distance  (Fig.  49);  distance  between  eyes  less  than  eye 

height  (Fig.  49)  [Nearctic]     Microdontomerus  anthidii  (Ashmead) 

Eye  height  2.5 X  or  less  than  malar  distance  (Fig.  50);  distance  between  eyes  equal  to 
eye  height  (Fig.  50)  [Nearctic]    Microdontomerus  apianus  Grissell 


Adontomerus  Nikol'skaya  postmarginal  vein,  2  to  5x  the  length  of  the 

stigmal  vein,  and  marginal  +  postmarginal 

Recognition. — Adontomerus  is  recognized      veins  equal  to  0.2 X  the  length  of  the  wing; 

by  the  straight  anterior  edge  of  the  meta-      the  occipital  carina  visible  in  dorsal  view  as 

pleuron  (Fig.  2);  the  fore  wing  with  mar-      a    finely    polished    line   raised    distinctly 

ginal   vein   1    to   2.5  X    the   length   of  the     above  surface  sculpture,  medially  arched 


240 


Journal  of  Hymenoptera  Research 


and  midway  between  hind  ocelli  and 
occipital  foramen  (Fig.  17);  and  the  hind 
femur  ventrally  without  a  tooth  (as  in 
Fig.  34). 

Number  of  Species. — 8. 

Number  Associated  with  Bees. — 2. 

Distribution. — Species  of  this  genus  are 
reported  in  the  Palearctic  Region  including 
the  former  Soviet  Union,  Bulgaria,  former 
Yugoslavia,  Hungary,  Italy,  Sardinia, 
Spain,  Jordan,  and  Algeria. 

Hosts  of  Genus. — Species  of  Adontomerus 
have  been  reared  from  cocoons  of  Lasio- 
campidae  (Lepidoptera),  galls  of  Cynipi- 
dae  (Hymenoptera),  and  cocoons  of  Mega- 
chiiidae  (Hymenoptera).  In  the  National 
Museum  of  Natural  History,  Washington, 
DC,  there  are  specimens  reared  from 
weevils  in  seed  heads  of  Asteraceae. 

Discussion. — Records  for  the  species 
listed  below  have  been  cited  in  the  litera- 
ture under  the  genus  Mellitotorymus ,  which 
was  synonymized  with  Adontomerus  by 
Grissell  (1995). 

Adontomerus  gregalis  (Steffan) 

Distribution.— PALEARCTIC:  Reported 
only  from  Sardinia  (Steffan  1964). 

Host. — Reared  from  Pseudoanthidium  (re- 
ported as  Anthidium)  lituratum  (Megachili- 
dae). 

Discussion. — I  believe  that  this  species 
and  Adontomerus  nesterovi  are  synonyms, 
but  I  have  not  seen  material  of  the  latter 
to  confirm  this.  Both  share  essentially 
similar  descriptions  as  well  as  the  same 
host.  I  treat  them  separately  here  to 
retain  the  known  data,  but  there  is  no 
way  to  distinguish  the  species  as  far  as  I 
can  tell. 

Adontomerus  nesterovi  Zerova 

Distribution.— PALEARCTIC:  Reported 
from  Turkmenistan  (Zerova  and  Roma- 
senko  1986). 

Host. — Reared  from  cocoons  of  Pseu- 
doanthidium (as  Paraantliidiellum)  lituratum 
(Megachilidae). 


Discussion. — Zerova  and  Romasenko 
(1986)  keyed  and  figured  this  species  in 
a  paper  on  the  parasitoids  of  megachilid 
bees  in  the  former  Soviet  Union. 

Echthrodape  Burks 

Recognition. — Echthrodape  is  recognized 
by  the  straight  anterior  edge  of  the  meta- 
pleuron  (as  in  Fig.  2)  and  by  the  relatively 
short  wing  venation  and  the  thickened 
marginal  vein  (Figs  9,  12,  13),  with  the 
postmarginal  vein  some  distance  from  the 
distal  edge  of  the  wing  (Fig.  9).  Additional 
characters  that  help  in  recognition  are  the 
toothed  hind  femur  (as  in  Fig.  38),  the 
developed  occipital  carina  that  lies  mid- 
way between  the  hind  ocelli  and  occipital 
foramen,  and  the  reduced  mouth  opening 
(Figs  14,  15,  indicated,  in  part,  by  the  long 
malar  distance)  with  reduced  mandibles 
(scarcely  visible  and  obscured  by  other 
mouth  parts). 

Number  of  Species. — 2. 

Number  Associated  with  Bees. — 2. 

Distribution. — The  genus  is  found  in  the 
Afrotropical  Region  in  Kenya,  and  in  the 
Australasian  Region  from  Papua  New 
Guinea. 

Host. — Hosts  for  both  species  belong  to 
the  genus  Braunsapis  (Apidae). 

Discussion. — The  species  of  this  genus 
are  uncommonly  encountered  and  are 
presently  the  only  indigenous  torymid 
bee  parasitoids  known  from  sub-saharan 
Africa  and  Australasia.  The  lack  of  records 
for  these  areas  is  probably  the  result  of 
a  paucity  of  collecting  and  rearing  both 
bees  and  parasitoids. 


Echthrodape  africana  Burks 


Re- 


Distribution.— AFROTROPICAL: 
ported  from  Kenya  (Burks  1969). 

Host. — Reared  from  nests  of  Braunsapis 
(as  Allodapula)  (Apidae)  as  reported  by 
Burks  (1969)  and  expounded  upon  by 
Michener  (1969)  who  reported  the  follow- 
ing host  records:  Braunsapis  simplicipes,  B. 
rolini,  and  B.  rufipes. 


Volume  16,  Number  2,  2007 


241 


Biology. — Larvae  of  E.  africana  are  exter- 
nal feeders  on  pupae  of  Braunsapis  (Mich- 
ener  1969).  One  parasitoid  was  seen  per 
host.  The  bee  is  a  progressive  feeder  which 
uses  burrows  in  the  pith  of  dead  Lantana 
stems.  It  moves  its  larvae  and  pupae  about 
and  does  not  distinguish  between  its  own 
progeny  and  those  of  E.  africana. 

Morphology. — Michener  (1969)  illustrated 
and  described  the  peculiar  larva  of  this 
species  as  well  as  the  pupa. 

Discussion. — The  two  known  species  are 
relatively  easily  identified  based  on  the 
distinctive  heads  (Figs  14,  15)  and  wing 
veins  (Figs  12,  13)  as  well  as  their  disjunct 
distributions. 

Eclitlirodape  papnana  Boucek 

Distribution.— AUSTRALASIAN:  Known 
from  Papua  New  Guinea  (Boucek  1988) 
and  Australia  (R.  Matthews,  per.  comm.). 

Host. — Reared  from  cells  of  Braunsapis 
unicolor  (Apidae)  nesting  in  bamboo  inter- 
nodes  (R.  Matthews,  per.  comm.). 

Discussion. — A  voucher  specimen  for  the 
Australian  record  was  kindly  placed  in  the 
U.  S.  National  Museum  collection  by 
Robert  Matthews. 

Microdontomerus  Crawford 

Recognition. — Microdontomerus  is  recog- 
nized by  the  straight  anterior  edge  of  the 
metapleuron  (as  in  Fig.  2),  the  simple  hind 
femur  (as  in  Fig.  34),  the  absence  of  an 
occipital  carina  (Fig.  16),  and  the  marginal 
vein  short,  1  to  2.5  X  the  length  of  the 
postmarginal  vein  and  2  to  5X  the  length  of 
the  stigmal  vein  (Figs  45,  46). 

Number  of  Species. — 22. 

Number  Associated  with  Bees. — 4. 

Distribution. — This  genus  is  transconti- 
nental in  the  Nearctic,  but  limited  in  other 
regions  of  the  world.  In  the  Palearctic  it  is 
found  in  Spain,  Italy,  Algeria,  and  Libya, 
and  in  the  Afrotropical  Region  it  is  found 
in  Senegal.  [The  genus  was  reported  in 
India  (see  Farooqi  1986,  David  et  al.  1990), 
but   this   is   probably   a   misidentification 


resulting  from  the  confusion  in  names  that 
existed  at  the  time.] 

Hosts. — Species  are  reported  from  mega- 
chilid  bees  and  cynipid  gall-formers  (Hy- 
menoptera),  tephritids  (Diptera),  buprestid 
eggs  and  curculionids  (Coleoptera),  mantid 
eggs  (Mantodea),  and  coleophorids,  geli- 
chiids,  lasiocampids,  and  tortricids  (Lepi- 
doptera).  At  least  one  Nearctic  species 
attacks  saturniid  eggs  (Lepidoptera).  Spe- 
cies have  also  been  documented  as  facul- 
tative hyperparasitoids  of  braconids  (Hy- 
menoptera)  (Grissell  2005). 

Microdontomerus  anthidii  (Ashmead) 

Distribution. — NEARCTIC:  This  species 
has  been  collected  in  southern  California, 
USA. 

Host. — Reared  from  Dianthidium  pudicum 
consimile  (as  Anthidium  consimile)  (Mega- 
chilidae). 

Discussion. — Microdontomerus  anthidii,  M. 
enigma,  and  M.  parkeri  are  difficult  to  distin- 
guish. Generally  M.  anthidii  is  smaller 
(2.3  mm  or  less)  with  a  shorter  ovipositor 
(less  than  1.2  X  hind  tibia),  whereas  M.  parkeri 
is  larger  (up  to  3.0  mm)  with  a  longer 
ovipositor  (more  than  2x  hind  tibia).  Micro- 
dontomerus enigma  is  about  the  size  of  M. 
anthidii,  but  with  the  longer  ovipositor  of  M. 
parkeri.  Microdontomerus  anthidii  is  fairly 
easily  separated  from  the  other  two,  however, 
based  on  discrete  morphological  differences 
in  the  fore  wing:  M.  anthidii  has  a  complete 
setal  row  along  the  upper  anterior  margin  of 
the  costal  cell  (absent  in  the  other  two  species) 
and  the  basal  cell  is  closed  (open  in  the  other 
two  species).  It  appears  that  while  all  three 
species  attack  megachilid  bees,  M.  anthidii  is 
usually  associated  with  species  of  the  tribe 
Anthidiini  that  create  nests  of  resin  and  sand 
grains,  whereas  M.  parkeri  and  M.  enigma  are 
associated  with  Osminiini  and  Megachilini 
that  make  stem  nests. 

Microdontomerus  apianus  Grissell 

Distribution. — NEARCTIC:  Known  from 
California,  USA. 


242 


Journal  of  Hymenoptera  Research 


Host. — Reared  from  Megachile  montivaga 
(Megachilidae). 

Discussion. — In  addition  to  characters 
given  in  the  key,  this  species  differs  from 
M.  anthidii  in  having  the  intermalar  dis- 
tance about  1.7x  the  malar  distance  (about 
2.5  X  in  M.  anthidii),  and  in  having  the 
ovipositor  sheaths  subequal  to  the  body 
length  and  2.0-3.0 X  as  long  as  the  hind 
tibia  (in  M.  anthidii  ovipositor  sheaths 
subequal  to  metasoma  and  usually  less 
than  1.5X  as  long  as  hind  tibia). 

Microdontomerus  enigma  Grissell 

Distribution.— NEARCTIC:  Known  only 
from  one  locality  in  Nevada,  USA. 

Hosts. — Reared  from  Hoplitis  bullifacies 
(Megachilidae). 

Discussion. — This  species  is  phenotypi- 
cally  nearly  identical  to  M.  parkcri.  Char- 
acters to  separate  the  two  are  given  in 
the  key.  Somewhat  more  difficult  to 
assess  is  that  in  M.  enigma  the  longest 
diameter  of  the  lateral  ocellus  is  less  than 
the  ocellocular  distance,  whereas  it  is 
subequal  to  or  greater  than  the  distance 
in  M.  parkeri. 

Microdontomerus  parkeri  Grissell 

Distribution.—  NEARCTIC:  Widespread 
in  the  western  and  southwestern  United 
States. 

Hosts. — Reared  from  Megachilidae:  Ash- 
meadiella  bigeloviae,  AshineadieUa  cubiceps, 
Ashmeadiella  gillettei,  AshineadieUa  rufipes, 
Hoplitis  bullifacies,  Hoplitis  palmarum,  Mega- 
chile brevis,  and  Osmia  marginata. 

Biologi/. — Microdontomerus  parkeri  is  a  gre- 
garious parasitoid  within  individual  bee 
cells.  The  number  of  individuals  ranged 
from  2  to  33  per  cell,  with  an  average  of 
about  8-9.  For  these  rearings  the  total 
number  of  M.  parkeri  specimens  was  229 
females  and  125  males  for  a  sex  ratio  of  1.8 
to  1.  Ten  of  these  rearings  contained  no 
males  (Grissell  2005). 

Discussion. — This  species  has  also  been 
reared  from  Ancistrocerus  sp.  and  Leptoclii- 


lus  sp.  (Vespidae:  Eumeninae).  It  is  the 
most  common  and  widespread  species  of 
Microdontomerus  attacking  bees. 

Motiodontomerus  Westwood 

Recognition. — Monodontomerus  is  recog- 
nized by  the  straight  anterior  edge  of  the 
metapleuron  (as  in  Fig.  2),  the  presence  of 
a  frenal  line  on  the  scutellum  (as  in  Fig.  8), 
the  hind  femur  with  a  single,  apicoventral 
tooth  (Figs  35,  37;  though  in  one  species 
this  tooth  is  poorly  defined,  Fig.  36),  and 
by  the  well  developed  occipital  carina 
which  is  nearly  horizontal  on  its  dorsal 
margin  and  closer  to  the  occipital  foramen 
than  to  the  hind  ocelli  (Fig.  18). 

Number  of  Species. — 32. 

Number  Associated  with  Bees. — 19. 

Distribution. — The  species  of  this  genus 
are  widespread  throughout  the  Holarctic, 
and  somewhat  less  common  in  the  Neo- 
tropical (Cuba,  Mexico,  Colombia,  Argen- 
tina) and  Oriental  (Sri  Lanka,  India,  Paki- 
stan) regions. 

Hosts. — Numerous  hosts  are  known  for 
this  genus  including  families  in  Diptera, 
Hymenoptera,  and  Lepidoptera.  The  pri- 
mary hosts  are  solitary  aculeate  bees  and 
wasps,  sawflies,  and  moths  (including 
their  tachinid  and  ichneumonid  parasi- 
toids).  An  authentic  record  of  Monodonto- 
merus (undetermined  species)  attacking 
social  vespids  {Mischocyt tarns;  Litte  1979) 
in  Arizona  occurs  in  the  literature,  but 
voucher  specimens  are  now  lost  (Litte,  /'// 
litt.).  Unfortunately,  some  species  of  Mono- 
dontomerus are  extremely  difficult  to  tell 
apart  and  as  a  consequence  there  have 
been  many  misidentifications  resulting  in 
incorrect  host  records  for  some  species.  For 
example,  Monodontomerus  aereus  Walker 
has  been  reported  from  Megachile  muraria 
(now  =  M.  pariet /;m)(Constantineanu  et  al. 
1956),  but  this  would  not  be  considered 
a  host  based  on  the  majority  of  records, 
which  are  from  Lepidoptera  (Grissell  2000, 
Noyes  2003).  Monodontomerus  vicicellae 
(Walker),  a  common  parasitoid  of  larval 


Volume  16,  Number  2,  2007 


243 


Lepidoptera  and  sawflies,  was  reported  to 
be  reared  from  an  ichneumonid  parasitoid 
in  the  nest  of  Megachile  "ramulorum  Rond." 
(Rondani  1877),  which  is  a  nomem  nudum. 
There  are  no  other  records  from  bees  for 
this  species  and  the  host  record  is  consid- 
ered to  be  incorrect.  Similarly,  Monodonto- 
merus  minor  (Ratzeberg),  also  a  parasitoid 
of  Lepidoptera  and  sawflies,  has  been 
reported  from  several  bees,  but  while  these 
records  appear  in  lists  (e.g.,  Herting  1977) 
they  apparently  have  no  basis  in  the 
primary  literature. 

Discussion. — In  the  following  section, 
summary  data  are  documented  in  Grissell 
(2000)  unless  otherwise  specified.  Identifi- 
cation is  often  more  easily  based  on  male 
characters.  Although  females  predominate 
in  reared  series,  species  have  gregarious 
larvae  and  some  males  are  almost  always 
present. 

Monodontomerus  acrostigmus  Grissell 

Distribution.— NEARCTIC:  Eastern  Tex- 
as, USA. 

Hosts. — Reared  from  pupa  of  Megachile 
sp.  (Megachilidae)  in  a  "mud-dauber 
nest". 

Discussion. — Monodontomerus  acrostigmus 
is  similar  in  appearance  to  M.  obscurus,  but 
differs  from  it  (and  all  other  known 
species)  by  having  the  stigma  posteriorly 
appendiculate  (Figs  51,  52).  In  addition,  it 
differs  from  M.  obscurus  by  having  the 
distal  portion  of  the  postmarginal  vein  one 
half  or  less  than  the  proximal  portion 
(subequal  in  M.  obscurus)  and  in  males, 
which  have  the  face  lateral  to  the  scrobal 
basin  distinctly  depressed  (not  depressed 
in  M.  obscurus). 

Monodontomerus  aeneus  (Fabricius) 

Distribution.— NEARCTIC:  Widespread 
throughout  the  northern  United  States 
and  southern  Canada.  PALEARCTIC:  Re- 
portedly widespread  in  western  Europe 
(Nikol'skaya  and  Zerova  1978)  and  often 
confused    with   M.    obscurus,   which    has 


the  same  distribution  and  general  host 
range. 

Hosts. — There  are  a  great  number  of 
hosts  listed  for  this  species  (as  obsoletus) 
in  the  Old  World  (see  Grissell  1995).  Only 
bee  hosts  are  listed  here  because  these  are 
certainly  correct  whereas  all  other  hosts  are 
suspect.  Old  World:  Anthophora  retusa, 
Ceratina  callosa  (Apidae);  Anthidium  floren- 
tinum,  Hoplitis  (as  Osmia)  adunca,  Megachile 
parietina  (as  Chalicodoma  muraria)  (and  Stelis 
nasuta,  a  cleptoparasite  of  this  host),  Mega- 
chile apicalis,  Megachile  centuncularis,  Mega- 
chile (as  Chalicodoma)  sicula,  Osmia  (as 
Metallinella)  brevicornis,  Osmia  coerulescens, 
Osmia  rufa  cornigera,  Osmia  cornuta,  Osmia 
emarginata,  Osmia  fulviventris,  Osmia  latreil- 
lei,  Osmia  rufa,  Osmia  submicans,  Osmia 
tricornis  (all  Megachilidae).  New  World: 
Verifiable  records  for  this  species  include 
Megachile  concinna,  Megachile  rotundata,  and 
Osmia  nigrifrons  (Megachilidae). 

Biology.—  Newport  (1849,  1852,  1853) 
provided  information  and  illustrations  of 
the  larvae,  their  digestive  tract,  and  feeding 
habits.  Johansen  and  Eves  (1966)  and  Eves 
(1970)  (and  possibly  Hobbs  and  Krunic 
1971)  published  biological  information  on 
this  species  (as  obscurus,  reidentified  by  me, 
based  upon  Eves'  specimens)  as  a  parasit- 
oid of  Megachile  rotundata.  Females  ovipos- 
ited through  the  leaf-lined  cell  and/or 
cocoon  of  the  host.  Between  3  and  51  eggs 
were  laid  externally  on  the  host.  An 
average  of  10  survived  in  one  study 
(Johansen  and  Eves  1966),  but  Bonelli  and 
Campadelli  (1990)  gave  an  average  of  24 
(range  =  10  to  51  adults  for  15  bee  cells). 
All  immature  stages  of  the  host  are 
vulnerable  to  attack  but  parasitization  of 
early  instars  is  rarely  successful.  Larvae  are 
non-cannibalistic.  The  life  cycle  can  be 
completed  in  about  20  days.  Goodpasture 
(1975)  detailed  the  mating  behavior  of  M. 
aeneus  (reported  as  M.  obscurus,  but  sub- 
sequently confirmed  as  M.  aeneus  in  Gris- 
sell 2000).  Tepedino  (1988a)  demonstrated 
that  7-12%  of  females  mated  before  emer- 
gence   from    the    host   cocoon.    He    also 


244 


Journal  of  Hymenoptera  Research 


showed  (Tepedino  1988b)  that  females  had 
an  initial  obligatory  requirement  for  host 
cocoon  and  prepupal  authenticity,  but  after 
24  hours  this  would  break  down  and 
females  would  oviposit  into  gelatin  cap- 
sules holding  bee  prepupae  or  even  agar 
replicates  of  bees.  Females  oviposited  onto 
fresh  host  prepupae  or  prepupae  that  were 
up  to  16  days  old.  Tepedino  (1988c) 
showed  that  superparasitism  occurs  but 
that  rates  go  down  as  resident  parasitoids 
become  older.  In  Spain,  rates  of  parasitism 
for  M.  aenens  (reported  as  M.  obsoletus)  on 
Osmia  cornuta  (Megachilidae)  varied  from 
0.5%  (Bosch  1994b)  to  73%  (Bosch  1994a). 
According  to  Bosch  (1993)  53-76%  of 
managed  bee  cocoons  were  parasitized 
when  paper  straws  containing  bee  cells 
were  extracted  from  their  nesting  blocks, 
but  cells  left  in  grooved  boards  were  left 
untouched.  In  the  Nearctic  this  parasitoid 
(as  M.  obscurus)  reportedly  replaced  the 
native  species  M.  montivagus  in  the  mid- 
1960's  as  the  most  important  parasitoid  of 
the  alfalfa  leafcutting  bee  in  North  Amer- 
ica, but  then  was  itself  replaced  by  a  pter- 
omalid  in  the  mid  1970's  (Eves  1982).  A 
paper  on  control  of  an  unknown  species  of 
Monodontomerus  in  Utah  by  Brindley  (1976) 
undoubtedly  refers  to  this  species. 

Morphology. — Goodpasture  illustrated 
the  karyotype  of  M.  obsoletus  (1975,  re- 
ported as  M.  obscurus  but  confirmed  as  M. 
aeneus  by  Grissell  2000).  The  chromosomes 
number  4  in  males,  8  in  females.  Good- 
pasture (1975)  illustrated  male  scapes,  and 
Walther  (1983)  illustrated  antennal  sensil- 
lae  of  this  species. 

Discussion. — This  species  was  introduced 
into  the  Nearctic  in  the  1930's  (Johansen 
and  Eves  1966),  but  it  was  misidentified  as 
M.  obscurus.  Its  correct  identity  as  M. 
obsoletus  was  reported  by  Tepedino  (1989) 
based  upon  my  identification.  The  name 
has  since  been  changed  to  M.  aeneus  by 
Graham  (1992)  who  studied  the  type 
material  of  the  species  involved.  Almost 
all  previously  published  host  records  (e.g., 
Peck  1969)  for  M.  obscurus  are  wrong  and 


most  should  now  refer  to  M.  aeneus.  Both 
M.  aeneus  and  M.  obscurus  are  common  and 
widespread  and  are  among  the  two  most 
difficult  species  of  the  genus  to  distinguish 
from  each  other.  This  is  disconcerting 
because  they  are  economically  important, 
have  both  been  introduced  into  the  New 
World  along  with  the  alfalfa  leafcutter  bee, 
and  have  been  confused  with  each  other 
since  their  introductions.  Only  the  appar- 
ent absence  of  sculpture  (though  faint 
coriaceous  sculpture  may  be  apparent  at 
some  angles  of  view)  on  the  median  frenal 
area  and  the  construction  of  the  frenal  apex 
offer  reliable  diagnostic  information  to 
separate  these  two  species,  but  even  this 
can  be  difficult  to  interpret  on  occasion.  An 
additional  character  that  may  sometimes 
help  to  define  these  two  taxa  is  found  in 
the  mesepimeron.  In  M.  aeneus  the  entire 
mesepimeron  is  essentially  smooth  (po- 
lished) except  for  some  slight  reticulation 
(or  carinae)  above  the  ventral  margin.  In  M. 
obscurus  the  ventral  1/5  of  the  mesepi- 
meron below  the  transepimeral  sulcus  is 
reticulate  and  the  anterior  1/3  is  alutac- 
eous  to  lightly  reticulate. 

Monodontomerus  anthidiorum  (Lucas) 

Distribution.—  PALEARCTIC:  Found 
only  in  Algeria. 

Host. — Reared  from  Rhodanthidiuiu  sticti- 
cum  (Megachilidae). 

Biology. — This  species  was  reared  from 
the  larva  of  its  host.  According  to  Lucas 
(1849)  the  bee  nested  in  empty  snail  shells 
(Helix  spp).  The  larvae  were  gregarious 
with  40-50  specimens  of  M.  anthidiorum 
found  in  each  shell. 

Discussion. — This  species  apparently  has 
not  been  collected  since  its  original  de- 
scription. In  both  sexes  this  is  one  of  the 
most  distinct  species  of  the  genus  based  on 
the  elongated  clypeus  (Figs  23,  24). 

Monodontomerus  argentinus  Brethes. 

Distribution.— NEOTROPICAL:  Costa 
Rica,  Panama,  Colombia,  and  Argentina. 


Volume  16,  Number  2,  2007 


245 


Hosts. — Reared  from  cells  of  Eufriesea 
nigrescens  (as  Euplusia  longipennis)  (Apidae) 
in  Colombia.  A  species  oi  Megachile  (Mega- 
chilidae)  also  serves  as  host. 

Biology. — Sakagami  and  Sturm  (1965) 
reported  that  this  species  developed  on 
the  pupal  stage. 

Discussion. — Monodontomerus  argentinus 
is  similar  to  M.  mexicanus  especially  in 
proportions  of  the  head  and  antenna  and  in 
details  of  the  wing,  hi  both  sexes  of  M. 
argentinus  the  median  area  of  the  frenum  is 
highly  polished,  whereas  in  M.  mexicanus 
the  median  frenal  area  is  longitudinally 
sculptured  similar  to  the  lateral  areas. 

Monodontomerus  bakeri  Gahan 

Distribution.—  NEARCTIC:  Colorado, 
Utah,  Idaho,  USA,  and  Alberta,  Canada. 

Hosts. — Megachile  pugnata,  Megachile  re- 
lativa,  Megachile  rotundata,  Osmia  coloraden- 
sis,  and  Osmia  texana  (Megachilidae). 

Discussion. — This  species  is  relatively 
uncommon,  but  large  numbers  were 
trapped  from  Megachile  rotundata  blocks 
as  a  nuisance  species  at  the  USDA  Bee 
Biology  and  Systematics  Laboratory  in 
Logan,  Utah  (pers.  obs.).  Monodofitomerus 
bakeri  is  unique  among  species  of  the  genus 
in  two  ways.  The  absence  of  a  malar  sulcus 
(Fig.  28),  or  its  expression  as  a  greatly 
curving,  indefinite  line  (Fig.  27  ),  is  atypical 
compared  to  the  straight,  well-defined 
sulcus  found  in  most  other  species  (e.g., 
Fig.  26).  Also,  the  bulging  lower  face 
(Fig.  27)  is  not  found  in  any  other  species, 
all  of  which  have  the  area  essentially  flat 
(as  in  Fig.  26). 

Monodontomerus  brevicrus  Grissell 

Distribution.— NEARCTIC:  California, 
USA. 

Hosts. — Reared  from  nests  of  Osmia 
ribifloris  (Megachilidae). 

Discussion. — Monodontomerus  brevicrus 
resembles  M.  dianthidii  in  having  metaso- 
mal  tergum  2  dorsally  sculptured,  but  it  is 
separated   as   follows:   Both   sexes   of  M. 


brevicrus  have  the  distal  portion  of  the 
postmarginal  vein  about  one-third  the 
length  of  the  proximal  portion  (Fig.  31) 
(about  equal  in  M.  dianthidii)  and  the  rim  of 
the  scutellum  apically  even  in  width  and 
not  projecting  (apically  widened  and  some- 
what projecting  in  M.  dianthidii).  In  females 
of  M.  brevicrus  metasomal  tergum  6  is 
obtuse  in  profile  (as  in  Fig.  33)  (acute  in 
M.  dianthidii,  as  in  Fig.  32)  The  males  of  M. 
brevicrus  are  unique  among  New  World 
males  in  modifications  found  in  the  fore 
leg  and  in  the  sunken  lower  face.  In  males 
the  fore  leg  is  reduced  (Fig.  40)  with  the 
tibia  shorter  in  length  than  the  femur  and 
ventrobasally  concave,  and  the  tarsomeres 
shortened  with  the  claw  length  equal  to 
tarsomeres  3  and  4  (fore  leg  unmodified  in 
other  species,  cf.  Fig.  39). 

Monodontomerus  dementi  Grissell 

Distribution.—  NEARCTIC:  Wyoming 
and  Colorado,  USA. 

Hosts. — Dianthidiinn  heterulkei  (Megachi- 
lidae) [also  reared  from  the  factitious  host 
Megachile  rotundata  (Megachilidae)  in  the 
laboratory]. 

Biologxj. — Clement  (1976)  found  this  spe- 
cies feeding  on  prepupae  in  cocoons  of  D. 
heterulkei.  Goodpasture  (1975)  described 
the  mating  behavior,  which  is  identical  to 
that  of  Monodontomerus  montivagus. 

Discussion. — Monodontomerus  dementi 
and  M.  laticoruis  are  similar  in  having  the 
anterior  half  of  the  upper  mesepimeral  area 
reticulately  sculptured  and  the  anterodor- 
sal  corner  with  diagonal  striations  extend- 
ing nearly  to  transepimeral  sulcus  (Fig.  53). 
They  differ  in  the  characters  outlined  in 
couplet  11  of  the  key. 

Monodontomerus  dianthidii  Gahan. 

Distribution. — NEARCTIC:  Eastern  Cali- 
fornia and  southwestern  Oregon,  USA. 

Hosts. — Dianthidiinn  sp.  (Megachilidae). 

Biology. — Reared  from  resin  nests. 

Discussion. — Monodontomerus  dianthidii  is 
phenetically  most  similar  to  A4.  brevicrus 


246 


Journal  of  Hymenoptera  Research 


based  upon  the  completely  sculptured 
frenal  area  and  metasomal  tergum  2; 
the  differences  between  these  species  are 
discussed  in  detail  under  M.  brevicrus 
above. 

Monodontomerus  laticornis  Grissell  and  Zerova 

Distribution.— PALEARCTIC:  Russia, 
Kazakhstan,  Ukraine,  and  Moldavia. 

Hosts. — Reared  from  Megachile  rotundata 
(Megachilidae);  Megachile  centuncularis  and 
Anthidium  florentinum  (Zerova  and  Stolbov 
1986)  (Megachilidae);  Anthidium  septemspi- 
nosum  (Zerova  and  Seryogina  (2002).  [A 
report  of  Apis  mellifera  as  host  (documented 
in  Noyes  2003)  seems  unlikely]. 

Biology. — This  is  a  gregarious  parasitoid 
within  cocoons  of  the  hosts. 

Discussion. — Zerova  and  Romasenko 
(1986)  key  and  figure  this  species  in  a  paper 
on  the  parasitoids  of  megachilid  bees  in  the 
Former  Soviet  Union.  This  species  and  M. 
clement i  are  similar  in  appearance,  and 
characters  to  distinguish  them  are  given 
under  couplet  11  of  the  key.  Monodo)ito- 
merus  laticornis  is  a  Palearctic  species  and 
M.  dementi  a  Nearctic  one,  so  they  should 
not  be  readily  confused. 

Monodontomerus  mandibularis  Gahan 

Distribution.— NEARCTIC:  Widespread 
throughout  the  eastern  USA  and  Canada 
from  Saskatchewan  south  to  Louisiana. 

Hosts. — Anthophora  abrupta,  A.  bomboides 
bomboides,  Melitoma  taurea  (Apidae);  Osmia 
cordata  (Megachilidae)  (Rau  1947). 

Biology. — Rau  (1947)  published  some 
preliminary  information  on  the  life  history 
of  this  species,  which  he  concluded  had 
one  or  two  generations  per  year.  He 
believed  the  wasp  to  be  a  primary,  gregar- 
ious parasitoid  of  its  host. 

Discussion. — Monodontomerus  mandibu- 
laris is  morphologically  similar  to  M. 
montivagus  but  differs  in  both  sexes  (and 
from  all  other  Monodontomerus  species)  by 
the  mandibles  having  a  single  large, 
ventral    tool       and    a    smaller,    subapical 


dorsal  one  (Fig.  48).  Other  species  have 
two  ventral  teeth  and  a  small  subapical 
dorsal  one  (as  in  Fig.  47)  or  have  the  dorsal 
tooth  so  reduced  as  to  be  easily  overlooked. 
The  mandibles  are  not  generally  exposed, 
however,  so  that  for  practical  purposes  M. 
mandibularis  is  best  distinguished  from  M. 
montivagus  as  follows:  In  females  the 
intermalar  distance  is  about  3x  the  malar 
distance  (about  2X  in  M.  montivagus;  this  is 
the  result  of  the  malar  distance  being 
relatively  shorter  in  M.  mandibularis  and 
the  face  less  produced  ventrally  below  the 
eyes,  cf.  Figs  47,  48)  and  the  posterior 
outline  of  metasomal  tergum  6  is  deeply 
concave  (shallow  in  M.  montivagus,  cf. 
Figs  32,  33);  in  males  the  scape  (Fig.  57)  is 
laterally  compressed  and  distinctly  C- 
shaped  in  profile  with  dorsal  and  ventral 
arches  asymmetrical  (in  M.  montivagus  the 
scape  is  dorsoventrally  compressed  and 
nearly  symmetrically  curved  in  profile, 
Fig.  56,  sometimes  greatly  so). 

Monodontomerus  mexicanus  Gahan 

Distribution.— NEARCTIC/NEOTROPI- 
CAL:  Spotty  distribution  in  Arizona,  north- 
central  Mexico,  and  western  Panama. 

Hosts. — Megachile  peruviana  (Megachili- 
dae) (Rau  1947);  Ancyloscelis  apiformis  (as 
armata)  (Torchio  1974)  and  Anthophora 
marginata  (Apidae)  (Herting  1977). 

Discussioji. — This  species  has  also  been 
reared  from  Trypoxylon  mexicanum  (Gahan 
1941),  T.  monteverde,  and  Passaloecus  (  = 
Polemistus)  pusillus  (Rau  1947)  (all  Crabro- 
nidae).  It  has  been  seen  walking  on  the 
surface  of  Trypoxylon  mud  nests  and 
drilling  with  its  ovipositor  through  the 
mud  walls  (Brockmann  in  litt.).  It  is  similar 
to  M.  argentinus  and  is  discussed  under 
that  species. 

Monodontomerus  montivagus  Ashmead 

Distribution.— NEARCTIC:  Widespread 
throughout  southern  Canada  and  USA. 
NEOTROPICAL:  Southern  Mexico  (Guer- 
rero). 


Volume  16,  Number  2,  2007 


247 


Hosts. — This  species  has  been  reared 
from  the  following  bees.  Apidae:  Antho- 
phorn  abrupta,  Anthophora  bomboides  bom- 
boid.es,  Anthophora  bomboides  neomexicana, 
Anthophora  linsleyi,  ? Anthophora  occidentalis, 
lAnthophora  vallorum,  Bombns  morrisoni, 
IMelissoides  sp.,  Xylocopa  tabaniformis  orpi- 
fex.  Megachilidae:  Anthidium  collectum,  An- 
thidium emarginatum,  Anthidiun  Imormo- 
num,  Anthidium  nest,  Ashmeadiella  Califor- 
nia!, Dianthidium  curvatum  sayi,  Dianthi- 
diinn  pudicum  pudicum,  Dianthidium 
pudicum  consimile,  Hoplitis  anthocopoides 
nest,  Megachile  centuncularis,  Megachile  re- 
lativa,  Megachile  rotundata,  Osmia  sp.  cocoon 
(in  Ttypoxylon  politum  nest  [Crabronidae]), 
Osmia  cordata,  Osmia  kincaidii,  Osmia  lati- 
sulcata,  Osmia  lignaria,  Osmia  ribifloris, 
Osmia  sanrafaelae,  Osmia  texana,  Stelis  de- 
pressa. 

Biology. — This  is  a  gregarious,  external 
parasitoid  of  aculeate  Hymenoptera.  Al- 
though there  are  numerous  references  to 
this  species  in  the  literature  (see  Peck  1963), 
most  of  these  are  simply  host  records 
without  biological  data.  A  few  papers  cited 
by  Peck  are  of  interest  and  are  cited  below. 
Davidson  (1893:  153)  stated  that  females  of 
M.  montivagns  deposited  10  to  20  eggs  in 
each  cell  of  Xylocopa  tabaniformis  orpifex  and 
that  some  broods  were  all  males  while 
others  were  all  females.  Hicks  (1926:  224) 
stated  that  M.  montivagns  was  parasitic 
both  on  Anthophora  occidentalis  and  its 
parasitoid  Oryttus  mirandus,  thus  acting  as 
a  primary  and  secondary  parasitoid. 
Mickel  (1928:  72-73)  reared  415  specimens, 
of  which  94%  were  females,  from  21  cells 
of  Anthophora  occidentalis.  He  found  no 
hyperparasitic  relationship  on  the  same 
bee  host  as  reported  by  Hicks  (1926). 
Linsley  and  MacSwain  (1942:  409-411)  also 
reported  montivagns  as  both  a  primary  and 
a  hyperparasitoid  on  Anthophora  linsleyi 
and  its  mutillid  parasitoid  Photopsis  auraria 
(now  =  Sphaeropthalma  itnicolor).  These 
authors  discussed  the  courtship  behavior 
of  montivagns  and  stated  that  its  larvae  fed 
on  the  prepupal  stage  of  the  bee.  They 


stated  that  only  one  cell  (of  9)  had  mixed 
sexes  of  this  parasitoid,  the  others  being 
either  female  (average  26  per  cell)  or  male 
(average  40  per  cell).  In  later  rearings, 
however,  MacSwain  (1958:  395)  found 
mixes  of  males  and  females  in  each  of  four 
cells  of  A.  occidentalis.  The  sex  ratio 
(males:females)  varied  from  1  to  12  to  1  to 
30.  Rau  (1922)  found  a  ratio  of  1  to  6. 
Goodpasture  (1975)  described  and  illus- 
trated the  courtship  behavior  of  M.  mon- 
tivagns. It  is  apparent  from  the  literature 
and  from  reared  specimens  that  M.  mon- 
tivagns is  parasitic  on  bees,  wasps,  and 
their  nest  associates.  New  and  old  nests  of 
aculeate  Hymenoptera  are  complex  sites  of 
diverse  taxa,  behaviorial  types,  and  suc- 
cessional  faunas.  Therefore,  our  biological 
knowledge  of  M.  montivagns  is  almost 
wholly  inadequate. 

Morphology. — Goodpasture  (1975)  de- 
scribed and  illustrated  the  male  scapes 
and  the  haploid  karyotype.  This  species 
has  6  chromosomes  in  males,  12  in  females. 

Discussion. — Females  of  M.  montivagus 
are  morphologically  similar  to  other  spe- 
cies reared  from  bees  (e.g.,  M.  parkeri,  M. 
tepedinoi,  M.  torchioi,  M.  mandibularis),  but 
males  differ  notably  in  morphology  of  the 
scape.  The  differences  between  M.  monti- 
vagns and  the  others  mentioned  are  dis- 
cussed under  each  of  these  species. 

Monodontomerus  obscurus  Westwood 

Distribution.— NEARCTIC:  Widespread 
from  coast  to  coast  in  the  United  States 
and  southeastern  Canada.  [Undoubtedly 
introduced  into  the  Nearctic  along  with  its 
host  the  alfalfa  leafcutting  bee.]  PALEARC- 
TIC:  Reportedly  widespread  in  western 
Europe  (Nikol'skaya  and  Zerova  1978) 
and  probably  often  confused  with  M. 
aenens  which  appears  to  be  sympatric. 
The  species  is  also  reported  from  the 
oriental  Region  (India). 

Hosts. — Hoplitis  (as  Osmia)  adunca,  Mega- 
chile argentata,  Megachile  centuncularis, 
Megachile    cephalotes,    Megachile  flavipes, 


248 


Journal  of  Hymenoptera  Research 


Megachile  lanata,  Megachile  parietina  (as 
Ckalicodoma  muraria),  Megachile  rotundata, 
M.  willughbiella,  Osmia  cordata,  Osmia  corni- 
frons,  Osmia  latreillei,  O.  lignaria,  Osmia 
ribfloris,  Osmia  rufa  rufa,  Osmia  rufa  corni- 
gera,  Osmia  sanrafaelae,  (Megachilidae);  An- 
thophora  plumipes,  Xylocopa  fenestrata  (Api- 
dae). 

Biology. — In  Spain,  M.  obscurus  is  consid- 
ered to  be  extremely  destructive  to  the 
alfalfa  leafcutting  bee  industry  and  chemi- 
cal methods  of  control  have  been  devised 
(Asensio  1982).  Krunic  and  Radovic  (1973) 
reported  that  M.  obscurus  can  go  through 
a  number  of  generations  without  diapause 
and  that  diapause  could  be  interrupted 
after  keeping  them  for  a  time  at  5  C. 

Morphology. — Radu  and  Botoc  (1968) 
illustrated  female  genitalia  in  detail.  Mac- 
Donald  and  Krunic  (1971)  illustrated  the 
somatic  chromosomes  for  M.  obscurus, 
which  number  6  in  males  and  12  in 
females.  (This  differs  from  M.  aeueus  and 
thus  strengthens  the  case  for  reproductive 
isolation  between  these  two  nearly  identi- 
cal species.)  Baker  et  al.  (1985)  described 
and  illustrated  the  last  instar  larva  and 
pupa  of  this  species  (adult  identity  con- 
firmed by  examination  of  voucher  speci- 
mens in  North  Carolina  State  University 
Insect  Collection). 

Discussion. — Zerova  and  Romasenko 
(1986)  key  and  figure  this  species  in  a  paper 
on  the  parasitoids  of  megachilid  bees  in  the 
former  Soviet  Union.  This  species  is  similar 
to  M.  aeueus  and  is  often  reared  from  the 
same  species  of  host  in  the  same  locality.  I 
discuss  the  two  species  more  fully  under 
M.  aeueus,  above. 

Monodontomerus  osmiae  Kamijo 

D/sfn7?Hfz'on.— PALEARCTIC:  Known 
from  Japan  and  the  Russian  Far  East  and 
introduced  into  the  Nearctic  (Grissell 
2003). 

Hosts. — Osmia  cornifrons,  Osmia  excavata, 
and  Osmia  taurus,  (Megachilidae)  (Kamijo 
1963,  1965). 


Biology. — Iwata  and  Tachikawa  (1966) 
reported  a  preponderance  of  females  for 
rearings  of  this  species  from  Osmia  taurus. 
From  61  cocoons  emerged  87  males  and 
726  females.  The  number  of  parasitoids  per 
host  (counted  for  4  cocoons  only)  varied 
from  14  to  26. 

Discussion. — Zerova  and  Romasenko 
(1986)  key  and  figure  this  species  in  a  paper 
on  the  parasitoids  of  megachilid  bees  in  the 
former  Soviet  Union.  Grissell  (2003)  illus- 
trated the  peculiar  male  head  of  this 
species  based  on  specimens  collected  in 
Silver  Spring,  Montgomery  County,  Mary- 
land, USA.  Monodontomerus  japonicus  Ash- 
mead  was  reported  from  Osmia  taurus,  but 
this  record  is  undoubtedly  a  misidentifica- 
tion  of  M.  osmiae  (Grissell  1995).  Males  of 
this  species  are  easily  identified  by  the 
peculiar  head  (Figs  21,  22).  Females  have 
the  discal  area  entirely  setose. 

Monodontomerus  parkeri  Grissell 

Distribution. — NEARCTIC:  Known  from 
widespread  localities  in  western  North 
America  stretching  from  Alberta,  Canada 
to  New  Mexico,  USA. 

Host. — Anthophora  occidentalis  (Apidae). 

Discussion. — Monodontomerus  parkeri  ap- 
pears most  similar  to  M.  tepedinoi  and  their 
separation  is  discussed  under  the  latter 
species.  This  species  is  also  easily  confused 
with  M.  montivagus.  Males  of  the  two 
species  may  be  readily  distinguished  by 
comparing  scapes:  in  M.  parkeri  the  scape  is 
laterally  compressed  and  asymmetrically 
bent  with  the  apex  enlarged  and  a  polished, 
depressed  area  on  its  outer  side  that 
continues  onto  the  ventral  surface  (some- 
what as  in  Fig.  57);  in  M.  montivagus  the 
scape  is  dorsoventrally  compressed,  sym- 
metrically bent,  and  has  the  polished  area 
completely  ventral  (Fig.  56).  Additionally, 
in  both  sexes  of  M.  parkeri,  the  apex  of  the 
costal  cell  dorsally  has  few  setae  (0  to  3  as 
in  Fig.  43)  whereas  in  M.  montivagus  there 
is  a  dorsal  row  of  setae  in  the  apical  half  to 
quarter  (as  in  Fig.  31),  and  the  transepim- 


Volume  16,  Number  2,  2007 


249 


eral  sulcus  is  complete  (Fig.  54),  whereas  in 
M.  montivagus  it  is  not  (Fig.  55).  There  are 
several  less  obvious  and  more  relative 
characters  that  are  difficult  to  use  without 
comparative  material.  In  M.  parkeri  the 
frenal  area  appears  medially  polished  un- 
der reflected  light  even  though  it  is 
sculptured,  whereas  in  M.  montivagus  this 
area  is  generally  entirely  sculptured.  In  M. 
parkeri  the  admarginal  wing  area  contains 
a  few,  widespaced  setae  (Fig.  44),  and 
relatively  few  setae  (3  to  5)  are  directly 
adjacent  to  the  marginal  vein  (so  that  there  is 
no  setal  row  parallel  to  the  vein),  but  in  M. 
montivagus  this  area  is  evenly  setose  to  the 
marginal  vein  (as  in  Fig.  43);  there  are  en- 
ough setae  to  form  a  row  parallel  to  the  vein. 

Monodontomerus  rugulosus  Thomson 

Distribution— PALEARCTIC:  Wide- 
spread in  western  and  central  Europe 
(Zerova  and  Seryogina  2002). 

Hosts. — This  species  has  been  reared 
from  Megachile  rotundata  (Megachilidae) 
(Zerova  and  Romasenko  1986). 

Biology. — This  is  a  gregarious  parasitoid 
in  cocoons  of  its  host. 

Discussion. — Monodontomerus  rugulosus 
appears  quite  similar  to  M.  argentinus,  but 
the  two  species  occur  in  different,  widely 
spaced  zoogeographic  regions.  They  may 
be  separated  by  characters  given  in  the  key. 

Monodontomerus  tepedinoi  Grissell 

Distribution. — NEARCTIC:  Known  from 
Oregon  and  Utah,  USA. 

Hosts. — The  species  has  been  reared 
from  Osmia  lignaria  (Megachilidae). 

Discussion. — Females  of  M.  tepedinoi  are 
easily  confused  with  M.  montivagus  and  M. 
parkeri.  From  M.  montivagus  it  is  most  readily 
separated  by  the  upper  anterior  margin  of 
the  costal  cell  with  only  1  to  3  setae  at  the 
apex  (as  in  Fig.  43),  whereas  in  M.  montiva- 
gus the  upper  anterior  margin  has  a  setal 
row  in  its  apical  1/4  to  1/3  (as  in  Fig.  31). 
From  M.  parkeri  it  is  separated  by  the  longer 
ovipositor  (ca.  1.5  to  almost  2X  the  metaso- 


mal  length;  1  to  1.2  X  in  M.  parkeri)  and  by 
the  scape,  which  has  some  metallic  green 
color  at  least  ventrally  (all  yellow  to  orange 
in  M.  parkeri).  Males  of  M.  tepedinoi  are  easier 
to  distinguish  than  females  based  on  the 
antenna  as  described  in  the  key  and 
compared  in  Figs  56,  57,  58).  Monodonto- 
merus tepedinoi  is  so  far  associated  only  with 
Megachilidae  and  M.  parkeri  with  Apidae. 

Monodontomerus  thorpi  Grissell 

Distribution. — NEARCTIC:  Known  from 
isolated  localities  in  southern  California, 
Arizona,  and  western  Texas,  USA. 

Hosts. — Reared  from  nests  of  Anthidium 
maculatum  (Megachilidae). 

Discussion. — This  species  has  been 
reared  from  twig  nests  in  the  eastern  and 
western  extremes  of  its  distribution.  It  is 
one  of  the  easiest  species  of  the  genus  to 
identify  in  both  sexes  as  it  is  the  only 
species  to  have  the  first  two  flagellar 
segments  reduced  (i.e.,  ring-like,  Fig.  19), 
whereas  all  other  species  have  only  the  first 
segment  reduced  (Fig.  20).  Additionally, 
the  hind  femur  is  enlarged  with  only 
a  ventral  angle  (Fig.  36)  as  opposed  to 
other  species  that  have  a  distinct  tooth 
(Figs  37,  38). 

Monodontomerus  torchioi  Grissell 

Distribution. — NEARCTIC:  The  species  is 
known  only  from  Utah,  USA. 

Hosts. — Reared  from  nests  of  Osmia 
lignaria  and  O.  sanrafaelae  (Megachilidae). 

Discussion. — Monodotitomerus  torchioi  is 
easily  confused  with  M.  montivagus,  M. 
tepedioni,  and  M.  parkeri  in  females.  The 
diagnostic  characters  used  to  separate 
these  three  taxa  are  given  in  key  couplets 
15  and  24-25  and  under  the  discussion  of 
the  species  mentioned. 

Pseudotorymus  Masi 

Recognition. — Anterior  margin  of  meta- 
pleuron  straight  (as  in  Fig.  2);  occipital 
carina  medially  arched  and  midway  be- 


250 


Journal  of  Hymenoptera  Research 


tween  the  hind  ocelli  and  occipital  foramen 
(as  in  Fig.  17);  hind  femur  ventrally  with 
a  slight  indication  of  a  tooth;  marginal  vein 
long,  3  to  7x  length  of  postmarginal  vein 
and  at  least  6X  length  of  stigmal  vein. 

Number  of  Species. — 43. 

Number  Associated  with  Bees. — 1  (ques- 
tionably). 

Distribution. — The  genus  is  most  abun- 
dant in  the  Palearctic  Region  (30  species) 
where  its  species  are  widespread  and 
extend  into  northern  Africa.  It  is  also 
known  from  the  Afrotropical  Region  (7 
species)  from  Madagascar,  Mali,  Mozam- 
bique, Nigeria,  Rwanda  (Republic  of  the 
Congo),  Senegal,  South  Africa,  and 
Sudan.  There  are  4  species  known  from 
India  in  the  Oriental  Region  and  a  single, 
widespread  species  is  known  from  the 
Nearctic  (southern  Canada  and  northern 
USA). 

Hosts. — Members  have  a  broad  host 
association  including  Curculionidae  (Co- 
leoptera)  in  leguminous  seed  pods;  Bruchi- 
dae  (Coleoptera)  from  galls  on  Asteraceae, 
Combretaceae,  Fabaceae,  Orchidaceae,  and 
Rubiaceae;  Cecidomyiidae  (Diptera)  asso- 
ciated with  Apiaceae,  Cruciferae,  Fabaceae, 
Lamiaceae,  Rosaceae,  Salicaceae,  and  Scro- 
phulariaceae;  Eurytomidae  (Hymenoptera) 
in  grass  stems  (Poaceae);  Cynipidae  (Hy- 
menoptera) in  pods  of  Papaveraceae;  Ten- 
thredinidae  (Hymenoptera);  and  Pyralidae 
(Lepidoptera). 

Discussion. — The  inclusion  of  this  genus 
in  relation  to  bee  hosts  is  highly  question- 
able and  is  based  upon  the  single  record 
for  P.  indicus  as  indicated  below.  Among 
the  other  42  known  species  of  Pseudotor- 
ymus  the  use  of  bees  is  unknown  so  this 
record  is  likely  to  be  incorrect. 

Pseudotorymus  indicus  (Marti) 

Distribution. — This  species  is  known  only 
from  southern  India  (Uttar  Pradesh,  Tamil 
Nadu)  (Mani  1989). 

Hosts. — The  type  series  was  reared  from 
"flower   bud    galls"    on    Dalbergia   sissoo 


(Fabaceae).  Mani  (1989)  listed  the  host  as 
a  "leafcutting  bee". 

Discussion. — In  light  of  the  original  rear- 
ing and  the  entire  host  range  given  above,  I 
am  inclined  to  dismiss  this  record  until  it 
can  be  reconfirmed. 

Torytnus  Walker 

Recognition. — This  genus  is  easily  recog- 
nized by  the  anterior  edge  of  the  meta- 
pleuron  (usually  its  upper  half)  projecting 
forward  as  a  lobe  into  the  mesepimeron 
which  is  subdivided  into  upper  and  lower 
sections,  the  lower  of  which  is  delimited  by 
an  anterior  groove  (Fig.  1,  compare  with 
Fig.  2,  arrows). 

Number  of  Species. — Approximately  375. 

Number  Associated  with  Bees. — 3. 

Distribution. — All  zoogeographic  regions 
except  Australia  where  it  was  apparently 
introduced  (Grissell  1995). 

Hosts  of  Genus. — Members  of  this  genus 
are  mostly  parasitoids  of  larvae  of  gall- 
forming  Diptera  and  Hymenoptera.  A  few 
have  been  reared  from  bees,  and  a  few  are 
phytophagous  in  seeds. 

Discussion. — Until  1998  the  species  that 
parasitized  bees  were  treated  as  the  genus 
Diomorus  Walker.  Graham  and  Gijswijt 
(1998)  synonymized  Diomorus  under  Tor- 
ymus,  dividing  its  members  into  several 
species  groups  of  the  latter. 

Ton/nuts  armatus  (Boheman) 

Distribution. — This  species  is  widespread 
in  the  Palearctic,  being  reported  from 
Europe  (Graham  and  Gijswijt  1998)  and 
Japan  (Kamijo  1979).  It  was  possibly  in- 
troduced into  Papua  New  Guinea  (Boucek 
1988). 

Hosts. — Kamijo  (1979)  reported  T.  arma- 
tus from  Ceratina  japojiica  (Apidae)  in  Rubus 
twigs  (Rosaceae)  in  Japan. 

Discussion. — This  species  has  reportedly 
been  reared  from  several  genera  of  Crab- 
ronidae,  including  Rliopalum  (Box  1920) 
and  Crossocerus  (Gijswijt  1974),  and  seems 
to  be  associated  with  wasps  and  bees  that 


Volume  16,  Number  2,  2007 


251 


nest  in  the  stems  of  Rubus  (Graham  and 
Gijswijt  1998).  It  is  the  most  distinct  of  the 
three  Torymus  species  known  from  bees, 
having  the  hind  coxa  dorsally  bare  and 
smooth,  and  the  propodeum  without  cari- 
nae. 

Toymus  cupreus  (Spinola) 

Distribution. — The  species  is  widespread 
in  the  Palearctic  (Nikol'skaya  and  Zerova 
1978),  mostly  in  the  "southern  parts  and 
middle  of  Europe"  and  reaching  into  the 
Netherlands  (Graham  and  Gijswijt  1998).  It 
is  reported  from  Burma  in  the  Oriental 
Region  (Mani  and  Kaul  1972). 

Host. — The  original  hosts  given  by  Spi- 
nola included  7  species  of  cynipid  galls, 
but  these  all  probably  housed  aculeate  bees 
or  wasps.  Mani  and  Kaul  (1972)  reported 
the  species  as  "widely  distributed  as  [a] 
parasitoid  of  Osmia  sp.  (Megachilidae)  and 
Sphecidae." 

Biology. — Enslin  (1922)  illustrated  and 
discussed  the  larval  and  pupal  stages  of 
this  species  (as  Diomorus  kollari). 

Discussion. — This  Palearctic  species  and 
the  following  Nearctic  species  are  geo- 
graphically separated  but  show  no  mor- 
phological differences.  In  coloration,  how- 
ever, they  are  distinct  as  explained  in  key 
couplet  3. 

Torymus  zabriskii  (Cresson) 

Distribution. — The  species  is  widespread 
in  the  United  States. 

Hosts. — The  only  reported  bee  host  is 
Ceratina  dupla  (Apidae)  (Zabriskei  1890). 

Biology. — Krombein  (1964)  reported 
some  short  biological  notes  on  this  species 
(as  Diomorus)  as  a  parasitoid  of  Ectemnius 
paucimaculatus  (Crabronidae).  He  sug- 
gested that  T.  zabriskii  parasitized  several 
cells  in  a  succession  of  cells  and  that 
oviposition  was  probably  through  the  wall 
of  the  plant  stem  {Hibiscus:  Malvaceae)  in 
which  the  wasp  nested. 

Discussion. — Ceratina,  the  only  reported 
bee  host  (Zabriskei  1890),  has  been  listed  in 


the  secondary  literature  several  times  but 
has  never  been  reconfirmed.  A  number  of 
other  hosts  in  the  family  Crabronidae  have 
been  reported  for  this  species  including 
Ectemnius,  Crossocerus,  and  Rhopalum  (sum- 
marized by  Grissell  1995). 

ACKNOWLEDGMENTS 

1  thank  Robert  Matthews,  University  of  Georgia, 
Athens,  for  supplying  unpublished  collecting  and 
rearing  information  for  Echthrodape  papuana  from 
Australia  and  for  donating  a  specimen  to  the  U.  S. 
National  Museum  of  Natural  History.  I  especially 
thank  Terry  Griswold,  USDA  Bee  Biology  and 
Systematica  Laboratory,  Logan,  Utah,  for  checking 
the  bee  names  used  in  this  paper.  His  help  has  been 
most  appreciated,  but  I  remain  responsible  for  errors 
in  their  subsequent  use.  For  reading  the  host  list  and 
offering  suggestions  on  host  data  I  am  grateful  to 
Jerome  Rozen,  American  Museum  of  Natural  History, 
New  York,  and  Frank  Parker  (retired),  USDA  Bee 
Biology  and  Systematics  Laboratory.  For  additional 
help  with  information  regarding  hosts  I  thank  John 
Brown,  Systematic  Entomology  Laboratory,  Washing- 
ton, DC.  For  reading  the  manuscript  and  offering 
positive  criticism  I  thank  Sam  Droege,  U.  S.  Geological 
Survey,  Laurel,  Maryland,  and  Norman  Woodley, 
Thomas  Henry,  and  David  Nickle,  Systematic  Ento- 
mology Laboratory.  Additionally,  the  editor  of  journal 
of  Hymenoptera  Research  and  several  reviewers  have 
provided  a  number  of  suggestions  and  corrections  to 
the  manuscript  for  which  I  am  grateful. 

LITERATURE  CITED 

Asensio,  E.  1982.  Leafcutter  bee  management  in  Spain: 
problems  of  parasitism.  Pp.  71-79  in:  G.  H.  Rank. 
Proceedings  of  the  first  international  symposium  on 
alfalfa  leafcutting  bee  management.  Saskatoon,  Sas- 
katchewan, Canada:  University  of  Saskatchewan 
Printing  Services.  281  pp. 

Baker,  J.  R.,  S.  B.  Kuhn,  and  E.  D.  Bambara.  1985.  Nests 
and  immature  stages  of  leafcutter  bees  (Hyme- 
noptera: Megachilidae).  Journal  of  the  Kansas 
Entomological  Society  58:  290-313. 

Baur,  H.  and  F.  Amiet.  2000.  The  Leucospidae 
(Hymenoptera:  Chalcidoidea)  of  Switzerland, 
with  a  key  and  data  on  the  European  species. 
Revue  Suisse  de  Zoologie  107:  359-388. 

Bonelli,  B.  and  G.  Campadelli.  1990.  Note  biologiche 
su  Chalicodoma  parietina  Geoffr.  (Hymenoptera- 
Megachilidae).  Bollettino  dell'Istituto  di  Entomolo- 
gia  "Guido  Grandi"  delta  Universita  degli  Studi  di 
Bologna  44:  1-9. 

Bosch,  J.  1993.  Parasitism  in  wild  and  managed 
populations  of  the  almond  pollinator  Osmia 
cornuta  Latr.  (Hymenoptera:  Megachilidae).  Jour- 


252 


Journal  of  Hymenoptera  Research 


nal   of  Agricultural   Research   31:   77-82.    (Journal 
dated  1992). 
— .  1994a.  Osmia  cornuta  Latr.  (Hym.,  Megachili- 


dae)  as  a  potential  pollinator  in  almond  orchards: 
Releasing  methods  and  nest-hole  length.  Journal  of 
Applied  Entomology  117:  151-157. 

1994b.  Improvement  of  field  management  of 


Osmia  cornuta  (Latreille)   (Hymenoptera,  Mega- 
chilidae)  to  pollinate  almond.  Apidologie  25:  71-83. 

Boucek,  Z.  1974.  A  revision  of  the  Leucospidae 
(Hymenoptera:  Chalcidoidea)  of  the  world.  Bul- 
letin of  the  British  Museum  (Natural  History), 
Entomology,  Supplement  23:  1-241. 

.  1988.  Australasian  Chalcidoidea  (Hymenoptera). 

Wallingford,  UK:  C.  A.  B.  International.  832  pp. 

Box,  L.  A.  1920.  [Note  with  exhibit  of  Hymenoptera]. 
Proceedings  of  the  South  London  Entomological  and 
Natural  History  Society  1919:  50. 

Brii.dley,  W.  A.  1976.  Carbaryl  control  of  chalcidoid 
parasitoids  from  alfalfa  leafcutting  bees.  Journal  of 
Economic  Entomology  69:  225-228. 

Burks,  B.  D.  1969.  New  Perilampidae  (Hymenoptera: 
Chalcidoidea).  Proceedings  of  the  Entomological 
Society  of  Washington  71:  73-81. 

Clement,  S.  L.  1976.  The  biology  of  Dianthidium 
heterulkei  heterulkei  Schwarz,  with  a  description 
of  the  larva  (Hymenoptera:  Megachilidae).  Was- 
mann  Journal  of  Biology  34:  9-22. 

Constantineanu,  M.  I.,  I.  Suciu,  I.  Andreiescu,  V. 
Ciochia,  and  C.  Pisica.  1956.  Contributii  la  studiul 
Chalcidoidelor  din  R.  P.  R.  Chalcidoide  (Chalci- 
doidea Ashmead  1899)  parazite  in  albilitia  pru- 
nului  (Aporia  crataegi  L.)  din  Moldova.  Analele 
Stiintifice  Universitatii  lasi  (New  Series)  2:  113-126. 

Davidson,  A.  1893.  The  nest  and  parasitoids  of 
Xylocopa  orpifex,  Smith  [sic].  Entomological  News 
4:151-153. 

David,  P.  M.  M.,  A.  M.  Hanifa,  and  S.  Natarajan.  1990. 
Biology  and  control  of  blossom  midge  Contarinia 
sp.  (Diptera:  Cecidomyiidae)  on  Jasminum  sambac 
in  Tamil  Nadu.  Entomon  15:  193-196. 

Enslin,  E.  1922.  Zur  Biologie  des  Solonius  rubicola  Duf. 
et  Perr,  (larvatus  Wesm.)  und  seiner  Parasiten. 
Konowia  1:  1-15. 

Eves,  J.  D.  1970.  Biology  of  Monodontomerus  obscurus 
Westwood  a  parasitoid  of  the  alfalfa  leafcutting 
bee,  Megachile  rotundata  (Fabricius)  (Hymenop- 
tera: Torymidae;  Megachilidae).  Melanderia  4: 
1-18. 

.  1982.  Evolution  of  predators,  nest  destroyers, 

and  parasitoids  of  the  alfalfa  leafcutting  bee  in  the 
Pacific  Northwest.  Pp.  57-59  in:  G.  H.  Rank. 
Proceedings  of  the  first  international  symposium  on 
alfalfa  leafcutting  bee  management.  Saskatoon,  Sas- 
katchewan, Canada:  University  of  Saskatchewan 
Printing  Services.  281  pp. 

Farooqi,  S.  I.  1986.  Family  Torymidae.  Pp.  259-277  in: 
B.  R.  Subba  Rao,  and  M.  Hayat,  eds.  The 
Chalcidoidea    (Insecta,    Hymenoptera)    of   India 


and  the  adjacent  countries.  Pt.  II.  A  Catalogue. 
Oriental  Insects  20:  1-430. 

Gahan,  A.  B.  1941.  A  revision  of  the  chalcid-flies  of  the 
genus  Monodontomerus  in  the  United  States 
National  Museum.  Proceedings  of  the  United  States 
National  Museum  90:  461-482. 

Gijswijt,  M.  J.  1974.  Faunistical  and  biological  records 
of  some  palaerctic  Chalcidoidea.  Entomologische 
Berichten  (Amsterdam)  34:  89-93. 

Goodpasture,  C.  E.  1975.  Comparative  courtship 
behavior  and  karyology  in  Monodontomerus  (Hy- 
menoptera: Torymidae).  Annals  of  the  Entomolog- 
ical Society  of  America  68:  391-397. 

Graham,  M.  W.  R.  1992.  Diagnostic  characters  of 
Monodontomerus  obscurus  Westwood,  1833  and  M. 
aeneus  (Fonscolombe,  1832)  (=  obsoletus  (Fabri- 
cius, 1798),  preocc.)  (Hym.,  Torymidae).  The 
Entomologist's  Monthly  Magazine  128:  143-146. 

and    M.   J.   Gijswijt.    1998.   Revision   of   the 

European  species  of  Torymus  Dalman  (s.  lat.) 
(Hymenoptera:  Torymidae).  Zoologische  Verhande- 
lingen  Leiden  317:  1-202. 

Grissell,  E.  E.  1995.  Toryminae  (Hymenoptera:  Chalci- 
doidea: Toryminae):  a  redefinition,  generic  clas- 
sification, and  annotated  world  catalog  of  species. 
Memoirs  on  Entomology,  International  2:  1-470. 
— .  2000.  A  Revision  of  New  World  Monodonto- 
merus Westwood  (Hymenoptera:  Chalcidoidea: 
Torymidae).  Contributions  of  the  American  Entomo- 
logical Institute  32  (1):  1-90. 

.   2003.   Discovery   of  Monodontomerus   osmiae 

Kamijo  (Hymenoptera:  Torymidae)  in  the  New 
World.  Proceedings  of  the  Entomological  Society  of 
Washington  105:  243-245. 

2005.  A  review  of  North  American  species  of 


Microdontomerus  Crawford  (Torymidae:  Hyme- 
noptera). Journal  of  Hymenoptera  Research  14: 
22-65. 

Herting,  B.  1977.  Hymenoptera.  A  catalogue  of  parasites 
and  predators  of  terrestrial  arthropods.  Section  A. 
Host  or  Prey/Enemy  4:  176.  Commonwealth  Agri- 
cultural Bureaux,  Institute  of  Biological  Control. 

Hesami,  S.,  M.  A.  Akrami,  and  H.  Baur.  2005.  Leucospis 
dorsigera  Fabricius  (Hymenoptera,  Leucospidae) 
as  a  hyperparasitoid  of  Cerambycidae  (Coleop- 
tera)  through  Xoridinae  (Hymenoptera:  Ichneu- 
monidae)  in  Iran.  Journal  of  Hymenoptera  Research 
14:  66-68. 

Hicks,  C.  H.  1926.  Nesting  habits  and  parasitoids  of 
certain  bees  of  Boulder  County,  Colorado.  The 
University  of  Colorado  Studies  (Series  A)  15: 
217-252. 

Hobbs,  G.  A.  and  M.  D.  Krunic.  1971.  Comparative 
behavior  of  three  chalcidoid  parasitoids  of  the 
alfalfa  leafcutter  bee,  Megachile  rotundata,  in  the 
laboratory.  The  Canadian  Entomologist  103: 
674-685. 

Iwata,  K.  and  T.  Tachikawa.  1966.  Biological  observa- 
tions on  53  species  of  the  superfamilies,  Chalci- 


Volume  16,  Number  2,  2007 


253 


doidea  and  Proctotrupoidea,  from  Japan  (Hyme- 
noptera:  Apocrita)  [sic].  Transactions  of  the  Shikoku 
Entomological  Society  9:  1-29. 

Johansen,  C.  and  J.  Eves.  1966.  parasitoids  and  nest 
destroyers  of  the  alfalfa  leafcutting  bee.  Washing- 
ton Agricultural  Experiment  Station,  Circular  469: 
1-12.' 

Kamijo,  K.  1963.  A  revision  of  the  species  of  the 
Monodontomerinae  occurring  in  Japan  (Hyme- 
noptera:  Chalcidoidea)  [Taxonomic  studies  on  the 
Torymidae  of  Japan,  II].  Insecta  Matsumurana  26: 
89-98. 

.  1965.  A  new  host-record  of  Monodontomerus 

osimae  Kamijo.  Insecta  Matsumurana  28:  78. 

1979.  Four  new  species  of  Torymidae  from 


Japan,  with  notes  on  two  known  species.  Akitu  (N. 
S.)  24:  1-11. 

Krombein,  K.  V.  1964.  Natural  History  of  Plummers 
Island,  Maryland.  XVIII.  The  hibiscus  wasp,  an 
abundant  rarity,  and  its  associates  (Hymenoptera: 
Sphecidae).  Proceedings  of  the  Biological  Society  of 
Washington  77:  73-112. 

Krunic,  M.  D.  and  I.  Radovic.  1973.  Some  aspects  of 
cold-hardiness  of  the  parasitoids  Monodontomerus 
obscurus,  Pteromalus  venustus  and  Melittobia  chaly- 
bii  (Chalcidoidea,  Hymenoptera)  and  their  host 
Megachile  rotundata  (Megachilidae,  Hymenop- 
tera). Arhiv  Bioloskih  Nauka,  Beograd  25:  45-48. 

Linsley,  E.  G.  and  J.  W.  MacSwain.  1942.  The 
parasitoids,  predators,  and  inquiline  associates 
of  Anthophora  linslei/i.  American  Midland  Naturalist 
27:  402-417. 

Litte,  M.  1979.  Mischocytarrus  flavitarsis  in  Arizona: 
social  and  nesting  biology  of  a  polistine  wasp. 
Zeitschrift  fiir  Rierpsychologie  50:  282-312. 

Lucas,  H.  1849.  Histoire  naturelle  des  animaux 
articules.  Insectes.  Exploration  Scientifique  de  l'AT 
gerie:  Sciences  Physiques  Zoologie.  Vol.  3.  Paris: 
Bertrand.  403  p. 

MacDonald,  M.  D.  and  M.  D.  Krunic.  1971.  Chromo- 
some numbers  of  Monodontomerus  obscurus  and 
Pteromalus  venestus,  chalcid  parasitoids  of  Mega- 
chile rotundata.  Arhiv  Bioloskih  Nauka,  Beograd  23: 
93. 

MacSwain,  J.  W.  1958.  Taxonomic  and  biological 
observation  on  the  genus  Hornia  (Coleoptera: 
Meloidae).  Annals  of  the  Entomological  Society  of 
America  51:  391-396. 

Mani,  M.  S.  1989.  The  fauna  of  India  and  adjacent 
countries.  Chalcidoidea  (Hymenoptera).  Pt.  I  and  II. 
Madras,  India:  Zoological  Survey  of  India.  1633  p. 

and    B.    K.    Kaul.    1972.    Some   Torymidae 

(Hymenoptera:  Chalcidoidea)  from  India.  Orien- 
tal Insects  6:  313-331. 

Michener,  C.  D.  1969.  Immature  stages  of  a  chalcidoid 
parasitoid  tended  by  allodapine  bees  (Hymenop- 
tera: Perilampidae  and  Apidae).  journal  of  the 
Kansas  Entomological  Society  42:  247-250. 


Michener,  D.  C.  2000.  The  bees  of  the  world.  The  Johns 
Hopkins  University  Press,  Baltimore 

Mickel,  C.  E.  1928.  The  biotic  factors  in  the  environ- 
mental resistance  of  Anthophora  occidentalis  Cress. 
(Hym.:  Apidae;  Dip.,  Coleop.).  Entomological  News 
39:  69-78. 

Newport,  G.  1849.  On  the  anatomy  and  development 
of  certain  Chalcididae  and  Ichneumonidae,  com- 
pared with  their  special  economy  and  instincts; 
with  descriptions  of  a  new  genus  and  species  of 
bee  parasitoid.  Part  I.  Proceedings  of  the  Linnean 
Society  of  Eondon  2:  23-26.  [November]. 

.    1852.    The   anatomy   and    development   of 

certain  Chalcididae  and  Ichneumonidae,  com- 
pared with  their  special  oeconomy  of  instincts; 
and  descriptions  of  a  new  genus  and  species  of 
bee  parasitoids.  The  Transactions  of  the  Linnean 
Society  of  London  21:  61-77.  [May  or  June]. 

.  1853.  Further  observations  on  the  habits  of 


Monodontomerus;  with  some  account  of  a  new 
Acarus  (Heteropus  ivntricosus),  a  parasitoid  in  the 
nests  of  Anthophora  retusa.  The  Transactions  of  the 
Linnean  Society  of  London  21:  95-102.  [June]. 

Nikol'skaya,  M.  N.  and  M.  D.  Zerova.  1978.  Family 
Torymidae  (Callimomidae)  (Torymids).  Pp. 
651-685  in:  G.  S.  Medvedev,  ed.  Keys  to  the 
Insects  of  the  European  part  of  the  USSR,  Vol.  3,  Part 
2.  New  Delhi  [Translated  from  the  Russian  by 
Amerind  Published  Co.,  1987].  1341  pp. 

Noyes,  J.  S.  1994.  The  reliability  of  published  host- 
parasitoid  records:  a  taxonomist's  view.  Nonve- 
gian  journal  of  Agricultural  Sciences  16:  59-69. 

.  2003.   Universal  Chalcidoidea  Database.  World 

Wide  Web  electronic  publication,  www.nhm.ac. 
uk/entomology/chalcidoids/ index.html  [ac- 
cessed 10-Sep-2003]. 

Peck,  O.  1963.  A  catalogue  of  the  Nearctic  Chalcidoi- 
dea. The  Canadian  Entomologist  Supplement  30: 
1-1092. 

.  1969.  Chalcidoid  parasitoids  of  the  alfalfa  leaf- 
cutter  bee,  Megachile  rotundata,  in  Canada.  The 
Canadian  Entomologist  101:  418-422. 

Radu,  V.  G.  and  M.  Botoc.  1968.  Contributii  la  studiul 
constitutiei  morfofunctionale  a  armaturii  genitale 
femele  la  calcidoide.  Babes-Bolyai,  Cluj  (Ser.  Biol.) 
13:  71-76. 

Rau,  P.  1922.  Ecological  and  behavior  notes  on 
Missouri  insects.  Transactions  of  the  Academy  of 
Science  of  St.  Louis  24:  1-71  pis.  5-8. 

.  1947.  Bionomics  of  Monodontomerus  mandibu- 

laris  Gahan,  with  notes  on  other  chalcids  of  the 
same  genus.  Antials  of  the  Entomological  Society  of 
America  40:  221-6. 

Rondani,  C.  1877.  Vesparia  parasita  non  vel  minus 
cognita.  Bollettino  della  Societa  Entomologica  Italiana 
9:  166-213. 

Rust,  R.  W.  1974.  The  systematics  and  biology  of  the 
genus  Osmia,  subgenera  Osmia,  Chalcosmia,  and 


254 


Journal  of  Hymenoptera  Research 


Cephalosmia  (Hymenoptera:  Megachilidae).  Was- 
mann  Journal  of  Biology  32:  1-93. 

Sakagami,  S.  F.  and  H.  Sturm.  1965.  Euplusia  long- 
ipeiuiis  (Friese)  und  ihre  merkwurdigen  brutzel- 
len  aus  Kolumbien.  Insecta  Matsumurana  28: 
83-92. 

Steffan,  J.  R.  1964.  Deux  nouveaux  genres  de  Torymi- 
dae  et  de  Cleonymidae  Palearctiques.  Entomo- 
phaga  9:  101-106. 

Stolbov,  N.  M.,  S.  M.  Palevych,  M.  D.  Zerova,  O.  F. 
Grobov,  T.  V.  Konovalov,  and  O.  P.  Shatskaya. 
1986.  [Recommendations  for  the  study  of  the 
parasitic  Chalcidoidea  of  Megachile  rotundata.]  [In 
Russian.]  Agricultural  Academy  of  the  USSR, 
Moscow 

Tepedino,  V.  J.  1988a.  Incidence  of  pre-emergence  sib- 
mating  in  Monodontomerus  obsoletus,  Pteromalus 
venustus,  and  Tetrastichus  megachilidis,  three  chal- 
cid  parasitoids  of  the  alfalfa  leafcutting  bee, 
Megachile  rotundata  (Hymenoptera:  Chalcididae). 
Pan-Pacific  Entomologist  64:  63-66. 

.  1988b.  Aspects  of  host  acceptance  by  Pteroma- 
lus venustus  Walker  and  Monodontomerus  obsoletus 
Fabricius,  parasitoids  of  Megachile  rotundata 
(Fabricius),  the  alfalfa  leafcutting  bee  (Hymenop- 
tera: Chalcididae).  Pan-Pacific  Entomologist  64: 
67-71. 

.  1988c.  Host  discrimination  in  Monodontomerus 

obsoletus  Fabricius  (Hymenoptera:  Torymidae), 
a  parasitoid  of  the  alfalfa  leafcutting  bee  Megachile 
rotundata  (Fabricius)  (Hymenoptera:  Megachili- 
dae). journal  of  the  New  York  Entomological  Society 
96:  113-118. 

.  1989.  Repulsion  of  three  chalcid  parasitoids  of 

the  alfalfa  leafcutting  bee,  Megachile  rotundata  (F.) 
by  the  fungicides  CAPTAN  and  DPX.  Pan-Pacific 
Entomologist  65:  403-409. 

Torchio,  P.  F.  1974.  Notes  on  the  biology  of  Ancyloscelis 
annata  Smith  and  comparisons  with  other  antho- 
phorine  bees  (Hymenoptera:  Apidae).  Journal  of 
the  Kansas  Entomological  Society  47:  54-62. 

Walther,  J.  R.  1983.  Antennal  patterns  of  sensilla  of 
the  Hymenoptera — a  complex  character  of  phy- 
logenetic  reconstruction.  Verhandlungen  der  Ver- 
eins  fiir  Naturzvissenschaftliche,  Hamburg  26: 
373-392. 

Zabriskei,  J.  L.  1890.  Society  notes.  Entomologica 
Americana  6:  100. 

Zerova,  M.  D.  and  L.  P.  Romasenko.  1986.  [Chalci- 
doids  (Hymenoptera,  Chalcidoidea),  parasitoids 
of  leafcutting  bees  (Hymenoptera,  Megachilidae), 
in  the  fauna  of  the  USSR].  [In  Russian].  Zoolo- 
gicheskii  Zhurnal  65:  1631-1643. 
-  and  L.  Ya.  Seryogina.  2002.  A  revision  of  Old 
World  Monodontomerus  (Hymenoptera:  Chalci- 
doidea: Torymidae).  National  Academy  of  Sciences 
of  Ukraine,  I.  I.  Schmalhausen  Institute  of  Zoology, 
Separate  Issue,  Kiev.  1-74. 


-  and  N.  M.  Stolbov.  1986.  New  data  on  the 
trophic  relation  of  Monodontomerus  laticornis 
Griss.  et  Zer.  (Hymenoptera,  Torymidae).  Vestnik 
Zoologii,  Kiev  1986  (5):  86-87. 


APPENDIX 

Two  lists  are  presented:  bee  host/chalci- 
doid  and  chalcidoid/bee  host.  The  bee  host 
list  presents  names  as  they  currently  are 
applied  in  the  literature  (i.e.,  valid  names), 
not  as  they  were  originally  published.  The 
lists  are  derived  from  Boucek  (1974),  Noyes 
(2003),  and  Grissell  (1995,  2000,  2005). 
Authors'  names  are  given  for  bee  host  in 
the  first  list;  chalcidoid  authors  are  given  in 
the  subsequent  list.  The  placement  of  bee 
genera  in  families  is  based  on  an  electronic 
version  (http://faculty.ucr.edu/~heraty/ 
beepage.html)  of  Michener  (2000). 

Bee  Host/Chalcidoid 

Apidae 

Allodape  exoloma  Strand:  Xylencyrtus  tridens 

Allodape  mucronata  Smith:  Xylencyrtus  tridens 

Allodape  panurgoid.es  Smith:  Xylencyrtus  tridens 

Allodape  rufogastra  Lepeletier  and  Serville:  Xy- 
lencyrtus tridens 

Allodapula  grandiceps  (Friese):  Xylencyrtus  tridens 

Allodapula  melanopus  (Cameron):  Xylencyrtus 
mumifex 

Ancyloscelis  apiformis  (F.):  Monodontomerus  mex- 
i can  us 

Anthophora  abrupta  Say:  Melittobia  acasta,  Melit- 
tobia  megachilis,  Pediobius  williamsoni,  Mono- 
dontomerus mandibular  is,  Monodontomerus 
montivagus 

Anthophora  bomboides  bomboides  Kirby:  Leucospis 
gigas,  Monodontomerus  mandibularis,  Monodon- 
tomerus montivagus 

Anthophora  bomboides  neomexicana  Cockerell: 
Monodontomerus  montivagus 

Anthophora  liusleyi  Timberlake:  Monodontomerus 
montivagus 

Anthophora  marginata  Smith:  Monodontomerus 
mexicanus 

Anthophora  occidentalis  Cresson:  Monodontomerus 
montivagus,  Monodontomerus  parkeri 

Anthophora  plumipes  (Pallas):  Monodontomerus 
obscurus 


Volume  16,  Number  2,  2007 


255 


Anthophora  retusa  (L.):  Melittobia  acasta,  Melitto- 
bia pelopoei,  Monodontomerus  aeneus 
1  Anthophora   vallorum    (Cockerell):   Monodonto- 
merus montivagus 
Apis  cerana  (¥.):  Antrocephalus  sp. 
Apis  mellifera  L.:  Dibrachys  boanniac,  Melittobia 
aeasta,  Monodontomerus  laticornis,  Nasonia  vi- 
tripennis,    Pteromalus   apum,   Tetrastichus   ho- 
wardi 
Bombus  agrorum  ¥.:  Pteromalus  conopidarum 
Bombus  amerieanorum  ¥:.  Pediobius  williamsoni 
Bombus  atratus  Franklin:  Pediobius  williamsoni 
Bombus  fervidus  F.:  Melittobia  chalybii 
Bombus  hortorum  (L.):  Melittobia  aeasta 
Bombus  lapidarius  (L.):  Pteromalus  conopidarum 
Bombus  morrisoni  Cresson:  Monodontomerus  mon- 
tivagus 
Bombus  ruderatus  (¥.):  Melittobia  aeasta 
Bombus  sp.:  Dibrachys  cavus,  Melittobia  haioaiien- 

sis,  Pachyerepoideus  vindemmiae 
Bombus  terrestris  (L.):  Melittobia  aeasta 
Braunsapis  leptozonia  (Vachal):   Xylencyrtus  tri- 

dens 
Braunsapis  rolini  (Vachal):  Echthrodape  africaua 
Braunsapis  rufipes  (Friese):  Echthrodape  africaua 
Braunsapis  simplicipes  Michener:  Echthrodape  af- 
ricaua 
Braunsapis  unicolor  Smith:  Echthrodape  papuana 
Centris  analis  ¥.:  Leucospis  cayennensis 
Centris  bicornuta  Mocsary:  Leucospis  cayennensis 
Centris  nitida  Smith:  Leucospis  cayennensis 
Centris  tarsata  (Smith):  Leucospis  cayennensis 
Centris  vittata  Lepeletier:  Leucospis  cayennensis 
Ceratina  calcarata  Robertson:  Axima  zabriskiei 
Ceratina  callosa  (¥.):  Eurytoma  nodularis,  Mono- 
dontomerus aeneus 
Ceratina  dallatorreana  Friese:  Eurytoma  apiculae 
Ceratina  dupla  Say:  Axima  zabriskiei,  Baryscapus 

amerieanus,  Habritys  latrus,  Torymus  zabriskii 
Ceratina  flavipes  Smith:  Neochalcis  breviceps 
Ceratina  ignara  Cresson:  Baryscapus  amerieanus 
Ceratina  japonica  Cockerell:  Cleonymus  ceratinae, 

Torymus  armatus 
Ceratina  nanula  Cockerell:  Baryscapus  amerieanus, 

Eurytoma  apiculae 
Ceratina  punctigena  Cockerell:  Eurytoma  apiculae 
Ceratina   sequoiac   Michener:    Baryscapus   ameri- 
eanus 
Ceratina    sp.:    Chciloneurus    leptulus,    Epistenia 
coeruleata,  Melittobia  megachilis,  Micrapion  da- 
lyi,  Micrapion  nasutum,  Micrapion  richardsi 
Ceratina  truncata  Friese:  Micrapion  steffaui 
ICtenoplectra  chalybea  Smith:  Leucospis  histrio 


Diadasina  distincta  (Holmberg):  Leucospis  genalis 

Eufriesea  nigrescens  (Friese):  Monodontomerus 
argentinus 

Euglossa  annectans  Dressier:  Melittobia  sp. 

Euglossa  ignita  Smith:  Polistomorpha  fasciata 

Euglossa  sp.:  Polistomorpha  couura,  Polistomorpha 
fasciata 

Eulaema  meriana  (Oliver):  Leucospis  pinna 

?Melissodes  sp.:  Monodontomerus  montivagus 

Melitoma  taurea  (Say):  Monodontomerus  mandibu- 
laris 

Trigona  sp.:  Brachymeria  discreta 

Xylocopa  brasilianorum  (L.):  Leucospis  klugii 

Xylocopa  caerulea  (¥.):  Coelopencyrtus  pallidiceps 

Xylocopa  caffra  (L.):  Coelopencyrtus  callainus, 
Coelopencyrtus  taylori 

Xylocopa  divisa  Klug:  Coelopencyrtus  callainus, 
Coelopencyrtus  taylori 

Xylocopa  fenestrata  (¥.):  Monodontomerus  obscurus 

Xylocopa  flavicollis  (De  Geer):  Coelopencyrtus 
callainus 

Xylocopa  flavorufa  (De  Geer):  Coelopencyrtus 
callainus,  Coelopencyrtus  taylori,  Coelopencyrtus 
watmoughi 

Xylocopa  frontalis  (Oliver):  Coelopencyrtus  gar- 
gar  is 

Xylocopa  inconstans  Smith:  Coelopencyrtus  callai- 
nus 

Xylocopa  nogueirai  Hurd  and  Moure:  Leucospis 
xylocopae 

Xylocopa  pubescens  Spinola:  Coelopencyrtus  sp. 

Xylocopa  scioensis  Gribodo:  Coelopencyrtus  cy- 
prius 

Xylocopa  sp.:  Leucospis  reversa 

Xylocopa  submordax  Cockerell:  Leucospis  anthi- 
dioides 

Xylocopa  sulcatipes  Maa:  Coelopencyrtus  sp. 

Xylocopa  tabaniformis  orpifex  Smith:  Monodonto- 
merus montivagus 

Xylocopa  tenuiscapa  Westwood:  Coelopencyrtus 
krishnamurtii 

Xylocopa  tranquebarorum  (Swederus):  Melittobia 
sosui 

Xylocopa  turanica  Morawitz:  Melittobia  acasta 

Xylocopa  watmoughi  Eardly:  Coelopencyrtus  sp. 

Colletidae 

Hylaeus  communis  Nylander:  Coelopencyrtus  are- 

narius,  Coelopencyrtus  callidii 
Hylaeus  cressoni  Cockerell:  Coelopencyrtus  hylaei 
Hylaeus  ellipticus  (Kirbv):  Coelopencyrtus  hylaeol- 

eter 
Hylaeus  fuscipennis  (Smith):  Coelopencyrtus  kaalae 


256 


Journal  of  Hymenoptera  Research 


Hylaeus  heraldicus  (Smith):  Coelopencyrtus  nothy- 
laei 

Hylaeus  koae  (Perkins):  Coelopencyrtus  kaalae 
Hylaeus  nigritus  (¥.):  Coelopencyrtus  arenarius 
Hylaeus  polifolii  Cockerell:  Eurytoma  stigmi 
Hylaeus  pubescens  (Perkins):  Coelopencyrtus  kaa- 
lae, Coelopencyrtus  sexramosus 
Hylaeus  sp.:  Eurytoma  nodularis,  Melittobia  acasta, 

Melittobia  hawaiiensis 
Hylaeus  varifrons  Cresson:  Pteromalus  analis 

Halictidae 

Halictus  africanus  Friese:  Aperilampus  varians 
ILasioglossum  pruinosum  (Robertson):  Eupelmus 

ashmeadi,  Eupelmus  rhizophelus 
Nomia  melauderi  Cockerell:  Mesopjolobus  brucho- 

phagi 

Megachilidae 

Anthidiellum  perplexum  Smith:  Leucospis  affinis 

Anthidiellum  sp.:  Leucospis  slossonae 

Anthidiellum  strigatum  (Panzer):  Leucospis  bifas- 
ciata,  Leucospis  dorsigera 

Anthidium  collectum  Huard:  Monodontomerus 
montivagus 

Anthidium  diadema  Latreille:  Leucospis  dorsigera 

Anthidium  emarginatum  (Say):  Leucospis  affinis, 
Leucospis  dorsigera,  Monodontomerus  montiva- 
gus 

Anthidium  florentinum  (F.):  Melittobia  acasta, 
Monodontomerus  aeneus,  Monodontomerus  lati- 
coruis 

Anthidium  maculatum  Smith:  Monodontomerus 
thorpi 

Anthidium  maculosum  Cresson:  Leucospis  affinis 

Anthidium  Imormonum  Cresson:  Monodonto- 
merus montivagus 

Anthidium  septemspi}iosum  Lepeletier:  Monodon- 
tomerus laticornis 

Ashmeadiella  aridula  astragali  Michener:  Leucospis 
affinis 

Ashmeadiella  bigeloviae  (Cockerell):  Microdonto- 
merus  parkeri 

Ashmeadiella  californica  (Ashmead):  Monodonto- 
merus montivagus 

Ashmeadiella  cubiceps  (Cresson):  Microdontomerus 
parkeri 

Ashmeadiella  gillettei  Titus:  Microdontomerus  par- 
keri 

Ashmeadiella  meliloti  Cockerell:  Leucospis  affinis 

Ashmeadiella  rufipes  Titus:  Microdontomerus  par- 
keri 


Coelioxys  octodentata  Say:  Aprostocetus  sp.,  Mer- 
isus  sp.,  Tetrastichus  coelioxydis 

ICoelioxys  quadridentatus  (L.):  Leucospis  gigas 

Dianthidium  curvatum  sayi  Cockerell:  Monodon- 
tomerus montivagus 

Dianthidium  heterulkei  Schwarz:  Monodontomerus 
dementi 

Dianthidium  pudicum  pudicum  (Cresson):  Leucos- 
pis affinis,  Monodontomerus  montivagus 

Dianthidium  pudicum  consimile  (Ashmead):  Mi- 
crodontomerus anthidii,  Monodontomerus  mon- 
tivagus 

Dianthidium  sp.:  Monodontomerus  dianthidii 

Heriades  crenulatus  Nylander:  Eurytoma  heriadi, 
Melittobia  acasta 

? Heriades  sp.:  Leucospis  dorsigera 

Heriades  truncorum  (L.):  Melittobia  acasta 

Hoplitis  acuticornis  (Dufour  and  Perris):  Leucospis 
biguetina 

Hoplitis  adunca  (Panzer):  Eurytoma  nodularis, 
Leucospis  dorsigera,  Melittobia  acasta,  Mono- 
dontomerus aeneus,  Monodontomerus  ob- 
scurus 

Hoplitis  anthocopoides  (Schenck)  (nest):  Monodon- 
tomerus montivagus 

Hoplitis  bullifacies  Michener:  Microdontomerus 
enigma,  Microdontomerus  parkeri 

Hopilitis  palmarum  (Cockerell):  Microdontomerus 
parkeri 

Hoplitis  producta  (Cresson):  Cleonymus  amabilis, 
Eurytoma  amplicoxa,  Eurytoma  stigmi,  Leucospis 
affinis 

Hoplitis  tridentata  (Dufour  and  Perris):  Leucospis 
biguetina,  Neochalcis  osmicida 

Hoplosmia  ligurica  (Morawitz):  Leucospis  dorsigera 

Lithurgus  capensis  Friese:  Leucospis  ornata,  Leu- 
cospis varicollis 

Megachile  aetheria  Mitchell:  Melittobia  hawaiiensis 

Megachile  albitarsis  Cresson:  Ablaxia  cupraeus 

Megachile  apicalis  Spinola:  Monodontomerus  ae- 
neus 

Megachile  argentata  (F.):  Dibrachys  cavus,  Melitto- 
bia acasta,  Monodontomerus  obscurus 

Megachile  bombycina  Radoszkowski:  Melittobia 
acasta 

Megachile  brevis  Say:  Aprostocetus  sp.,  Cricellius 
megachilis,  Leucospis  affinis,  Melittobia  chalybii, 
Merisus  sp.,  Microdontomerus  parkeri,  Tetrasti- 
chus coelioxydis 

Megachile  centuncularis  (L.):  Ablaxia  cupraeus, 
Anagrus  putnamii,  Aprostocetus  pygmaeus,  Bar- 
yscapus  megachilidis,  Dibrachys  sp.,  Melittobia 
acasta,  Melittobia  chalybii,  Melittobia  megachilis, 


1  ■  ' 


Volume  16,  Number  2,  2007 


257 


Monodontomerus  aeneus,  Monodontomerus  lati- 
cornis,  Monodontomerus  montivagus,  Monodon- 
tomerus obscurus,  Pteromalus  apum,  Pteromalus 
macronychivorus 

Megachile  cephalotes  Smith:  Monodontomerus  ob- 
scurus 

Megachile  concinna  Smith:  Baryscapus  megachili- 
dis,  Melittobia  australica,  Monodontomerus  ae- 
neus 

Megachile  disjunctiformis  Cockerell:  Leucospis 
japonica 

Megachile  ericetorum  Lepeletier:  Leucospis  dorsi- 
gera 

Megachile  flavipes  Spinola:  Monodontomerus  ob- 
scurus 

Megachile  gentilis  Cresson:  Baryscapus  megachili- 
dis,  Leucospis  affinis 

Megachile  gomphrenae  Holmberg:  Melittobia  ha- 
waiiensis 

Megachile  gratiosa  Cameron:  Melittobia  sp. 

Megachile  hungarica  Gerstaecker:  Leucospis  gigas 

Megachile  biennis  Provancher:  Leucospis  affinis, 
Melittobia  chah/bii 

Megachile  lanata  (¥.):  Melittobia  australica,  Mono- 
dontomerus obscurus 

Megachile  mendica  Cresson:  Leucospis  affinis 

Megachile  montivaga  Cresson:  Leucospis  affinis, 
Microdontomerus  apianus 

Megachile  nipponica  Cockerell:  Leucospis  japonica 

Megachile  pallefacta  Vachal:  Melittobia  hawaiiensis 

Megachile  palmarum  Perkins:  Melittobia  hawaiien- 
sis 

Megachile  parietina  (Geoffrey):  Leucospis  gigas, 
Melittobia  acasta,  Monodontomerus  aeneus, 
Monodontomerus  obscurus 

Megachile  peruviana  Smith:  Monodontomerus  mex- 
icanus 

Megachile  poeyi  Guerin-Meneville:  Leucospis  poeyi 

Megachile  pugnata  Say:  Dibrachys  sp.,  Leucospis 
affinis,  Melittobia  sp.,  Monodontomerus  bakeri 

Megachile  pyrenaica  Lepeletier:  Leucospis  gigas, 
Pteromalus  apum 

Megachile  rancaguensis  Friese:  Leucospis  hopei 

Megachile  rangii  Cheesman:  Leucospis  amino 

Megachile  relativa  Cresson:  Dibrachys  relativus, 
Leucospis  affinis,  Melittobia  acasta,  Melittobia 
chalybii,  Monodontomerus  bakeri,  Monodonto- 
merus montivagus,  Pteromalus  apum 

Megachile  rotundata  (F.):  Baryscapus  daira,  Bar- 
yscapus megachilidis,  Dibrachys  confusus,  Dibra- 
chys maculipenuis,  Melittobia  acasta,  Melittobia 
australica,  Melittobia  chalybii,  Melittobia  ha- 
waiiensis, Monodontomerus  aeneus,  Monodonto- 


merus bakeri,  Monodontomerus  dementi  (in  lab), 
Monodontomerus  laticornis,  Monodontomerus 
montivagus,  Monodontomerus  obscurus,  Mono- 
dontomerus rugulosus,  Pteromalus  apum,  Pter- 
omalus conopidarum,  Pteromalus  veneris,  Tetra- 
stichus  sp. 

Megachile  sculpturalis  Smith:  Leucospis  japonica 

Megachile  sicula  Rossi:  Leucospis  gigas,  Monodon- 
tomerus aeneus 

Megachile  sp.:  Brachymeria  paraguayensis,  Calosota 
fumipennis,  Horismenus  albipes,  Kocourekia  cla- 
vigera,  Leucospis  histrio,  Leucospis  intermedia, 
Melittobia  pelopoei,  Monodontomerus  acrostig- 
mus,  Monodontomerus  argentinus 

Megachile  spissula  (Cockerell):  Lariophagus  obtu- 
sus,  Melittobia  acasta 

Megachile  ustulatum  (Smith):  Leucospis  histrio 

Megachile  willozomorensis  Brauns:  Leucospis  ornata 

Megachile  willughbiella  (Kirby):  Melittobia  acasta, 
Monodontomerus  obscurus,  Pteromalus  apum 

Megachile  xylocopoides  Smith:  Baryscapus  mega- 
chilidis 

Megachile  zaptlana  Cresson:  Melittobia  australica 

Microthurge  corumbae  (Cockerell):  Leucospis  sp. 

Osmia  atriventris  Cresson:  Leucospis  affinis 

Osmia  bicolor  (Schrank):  Eulophus  osmiarum 

Osmia  bicornis  (Schrank):  Leucospis  dorsigera, 
Leucospis  gigas 

Osmia  brevicomis  (F.):  Monodontomerus  aeneus 

Osmia  californica  Cresson:  Leucospis  affinis 

Osmia  "coerulea"  [?lapsus  for  O.  coerulescens, 
Baur  and  Amiet  2000]:  Leucospis  gigas 

Osmia  coendescens  (L.):  Aprostocetus  pygmaeus, 
Eurytoma  nodularis,  Leucospis  gigas,  Monodon- 
tomerus aeneus 

Osmia  coloradensis  Cresson:  Monodontomerus 
bakeri 

Osmia  cordata  Robertson:  Monodontomerus  man- 
dibularis,  Monodontomerus  nuvitivagus,  Mono- 
dontomerus obscurus 

Osmia  cornifrons  Radoszkowski:  Monodonto- 
merus obscurus,  Monodontomerus  osmiae 

Osmia  cornuta  (Latreille):  Leucospis  dorsigera, 
Monodontomerus  aeneus 

Osmia  emarginata  Lepeletier:  Leucospis  interme- 
dia, Monodontomerus  aeneus 

Osmia  excavata  Alfken:  Leucospis  japonica,  Mono- 
dontomerus osmaie 

Osmia  fedtschenkoi  (Morawitz):  Leucospis  dorsi- 
gera 

Osmia  fulviventris  (Panzer):  Leucospis  dorsigera, 
Monodontomerus  aeneus 

Osmia  globicola  (Stadelmann):  Leucospis  osmiae 


258 


Journal  of  Hymenoptera  Research 


Osmia  kincaidii  Cockerell:  Leucospis  affinis,  Mono- 

dontomerus  montivagus 
Osmia   latisulcata   Michener:   Monodontomerus 

montivagus 

Osmia  latreillei  (Spinola):  Calosota  vernalis,  Mono- 
dontomerus aeneus,  Monodontomerus  obscurus 

Osmia  leueomelana  (Kirby):  Eurytoma  sp.,  Melit- 
tobia  acasta 

Osmia  lignaria  Say:  Monodontomerus  montivagus, 
Monodontomerus  obscurus,  Monodontomerus  te- 
pedinoi,  Monodontomerus  torchioi 

Osmia  marginata  Michener:  Microdontomerus 
parkeri 

Osmia  nigrifrons  Cresson:  Dibrachys  pelos,  Mono- 
dontomerus aeneus 

Osmia  niveata  (F.):  Leucospis  dorsigera 

Osmia  parietina  Curtis:  Leucospis  dorsigera 

Osmia  parvula  Dufour  and  Perris:  Eurytoma 
nodularis 

Osmia  pumila  Cresson:  Leucospis  affinis 

Osmia  ribifloris  Cockerell:  Monodontomerus  brevi- 
crus,  Monodontomerus  montivagus,  Monodonto- 
merus obscurus 

Osmia  rostrata  Sandhouse:  Leucospis  affinis 

Osmia  rufa  comigera  (Rossi):  Monodontomerus 
aeneus,  Monodontomerus  obscurus 

Osmia  rufa  rufa  (L):  Leucospis  dorsigera,  Leucospis 
gigas,  Melittobia  acasta,  Monodontomerus  ae- 
neus, Monodontomerus  obscurus 

Osmia  sanrafaelae  Parker:  Monodontomerus  mon- 
tivagus, Monodontomerus  obscurus,  Monodonto- 
merus torchioi 

Osmia  simiUima  Smith:  Leucospis  affinis 

Osmia  sp.:  Epistenia  coeruleata,  Monodontomerus 
montivagus  (cocoon  in  Trypargilum  politum 
nest),  Torymus  cupreus 

Osmia  submicans  Morawitz:  Monodontomerus 
aeneus 

Osmia  taurus  Smith:  Leucospis  japonica,  Mono- 
dontomerus osmiae 

Osmia  texana  Cresson:  Monodontomerus  bakeri, 
Monodontomerus  montivagus 

Osmia  tricornis  Latreille:  Leucospis  dorsigera, 
Monodontomerus  aeneus 

Pachyanthidium  cordatum  (Smith):  Leucospis  tri- 
color 

Pachyanthidium  truncataum  (Smith):  Leucospis 
tricolor 

Pseudoantliidium  lituratum  (Panzer):  Adontomerus 
gregalis,  Adontomerus  nesterovi,  Neochalcis  fer- 
toni 
Khodanthidium   sticticum    (F.):   Monodontomerus 
anthidiorum 


Serapista  denticulata  (Smith):  Leucospis  africana, 
Leucospis  tricolor 

Stelis  depressa  Timberlake:  Monodontomerus  mon- 
tivagus 

Stelis  nasuta  Latreille:  Melittobia  acasta,  Mono- 
dontomerus aeneus 

Stelis  sexmaculata  Ashmead:  Cleonymus  amabilis, 
Leucospis  affinis 

Chalcidoid/Bee  Host 

Chalcididae 

Antrocephalus  sp.:  Apis  ceraua 
Brachymeria  discreta  Gahan:  Trigona  sp. 
Brachymeria  paraguayensis  Girault:  Megachile  sp. 
Neochalcis  breviceps  (Masi):  Ceratina  fiavipes 
Neochalcis  fertoni  (Kieffer):  Pseudoanthidium  litur- 
atum 
Neochalcis  osmicida  (Saunders):  Hoplitis  tridentata 

Encyrtidae 

Cheiloneurus   leptulus    Annecke    and    Prinsloo: 

Ceratina  sp. 

Coelopencyrtus  arenarius  (Erdos):  Hylaeus  commu- 
nis, Hylaeus  uigritus 

Coelopencyrtus  callainus  Annecke:  Xylocopa  caffra, 
Xylocopa  divisa,  Xylocopa  flavicollis,  Xylocopa 
flavorufa,  Xylocopa  inconstans 

Coelopencyrtus  callidii  (Jansson):  Hylaeus  commu- 
nis 

Coelopencyrtus  cyprius  Annecke:  Xylocopa  scioen- 
sis 

Coelopencyrtus  gargaris  (Walker):  Xylocopa  fronta- 
lis 

Coelopencyrtus  hylaei  Burks:  Hylaeus  cressoni 

Coelopencyrtus  hylaeoleter  Burks:  Hylaeus  ellipti- 
cus 

Coelopencyrtus  kaalae  (Ashmead):  Hylaeus  fusci- 
pennis,  Hylaeus  koae,  Hylaeus  pubescens 

Coelopencyrtus  krishnamurtii  (Mahdihassan):  Xy- 
locopa tenuiscapa 

Coelopencyrtus  nothylaei  Annecke:  Hylaeus  heral- 
dicus 

Coelopencyrtus  pallidiceps  (Girault):  Xylocopa  caer- 
ulea 

Coelopencyrtus  sexramosus  Timberlake:  Hylaeus 
pubescens 

Coelopencyrtus  sp.:  Xylocopa  pubescens,  Xylocopa 
sulcatipes,  Xylocopa  watmouglii 

Coelopencyrtus  tai/lori  Annecke  and  Doutt:  Xylo- 
copa caffra,  Xylocopa  divisa,  Xylocopa  flavorufa 


Volume  16,  Number  2,  2007 


259 


Coelopencyrtus  watmoughi  Annecke:  Xylocopa 
flavorufa 

Eulophidae 

Aprostocetus  pygmaeus  Zetterstedt:  Megachile 
centuncularis,  Osmia  coerulescens 

Aprostocetus  sp.:  Coelioxys  octodeutata,  Megachile 
brevis 

Baryscapus  americanus  (Ashmead):  Ceratina  du- 
plet, Ceratina  ignara,  Ceratina  nanula,  Ceratina 
sequoiae 

Baryscapus  daira  (Walker):  Megachile  rotumiata 

Baryscapus  megachilidis  (Burks):  Megachile  cen- 
tuncularis, Megachile  concinna,  Megachile  genti- 
lis,  Megachile  rotundata,  Megachile  xylocopoides 

Eulophus  osmiarum  Robineau-Desvoidy:  Osmia 
bicolor 

Horismenus  albipes  (Schrottky):  Megachile  sp. 

Kocourekia  clavigera  Boucek:  Megachile  sp. 

Melittobia  acasta  (Walker):  Anthidiumflorentinuiu, 
Anthophora  abrupta,  Anthophora  retusa,  Apis 
mellifera,  Bombus  hortorwn,  Bombus  ruderatus, 
Bombus  terrestris,  Heriades  crenulatus,  Heriades 
truncorum,  Hoplitis  adunca,  Hylaeus  sp.,  Mega- 
chile argentata,  Megachile  bombycina,  Megachile 
centuncularis,  Megachile  parietina,  Megachile 
relativa,  Megachile  rotundata,  Megachile  spis- 
sula,  Megachile  willughbiella,  Osmia  leucome- 
lana,  Osmia  rufa,  Stelis  nasuta,  Xylocopa  tur- 
anica 

Melittobia  australica  Girault:  Megachile  concinna, 
Megachile  lanata,  Megachile  rotundata,  Mega- 
chile zaptlana 

Melittobia  chalybii  Ashmead:  Bombus  fervidus, 
Megachile  brevis,  Megachile  centuncularis, 
Megachile  inermis,  Megachile  relativa,  Megachile 
rotundata 

Melittobia  hawaiiensis  Perkins:  Bombus  sp.,  Hy- 
laeus sp.,  Megachile  aetheria,  Megachile  gom- 
phrenae,  Megachile  pallefacta,  Megachile  pal- 
marum,  Megachile  rotundata 

Melittobia  megachilis  (Packard):  Anthophora 
abrupta,  Ceratina  sp.,  Megachile  centuncularis 

Melittobia  pelopoei  [unavailable  name]:  Antho- 
phora retusa,  Megachile  sp. 

Melittobia  sosui  Dahms:  Xylocopa  tranquebarorum 

Melittobia    sp.:    Euglossa   annectans,   Megachile 

gratiosa,  Megachile  pugnata 
Pediobius  williamsoni  Girault:  Anthophora  abrupta, 

Bombus  americanorum,  Bombus  atratus 
Tetrastichus  coelioxydis  (Burks):  Coelioxys  octodeu- 
tata, Megachile  brevis 
Tetrastichus  howardi  (Olliff):  Apis  mellifera 


Tetrastichus  sp.:  Megachile  rotundata 
Torymus  armatus  (Boheman):  Ceratina  japonica 
Torymus  cupreus  (Spinola):  Osmia  sp. 
Torymus  zabriskii  (Cresson):  Ceratina  dupla 
Xylencyrtus  mumifex  Annecke:  Allodapula  mela- 

nopus 
Xylencyrtus  tridens  Annecke:  Allodape  exoloma, 
Allodape  mucronata,  Allodape  panurgoides,  Allo- 
dape rufogastra,  Allodapula  grandiceps,  Braunsa- 
pis  leptozonia 

Eupelmidae 

Calosota  fumipennis  Curtis:  Megachile  sp. 

Calosota  vernalis  Curtis:  Osmia  latreillei 

Eupelmus  ashmeadi  Melander  and  Brues:  ILasio- 
glossum  pruinosum 

Eupelmus  rhizophelus  Brues:  ILasioglossum  prui- 
nosum 

Eurytomidae 

Axima   zabriskiei   Howard:    Ceratina    calcarata, 

Ceratina  dupla 
Eurytoma  amplicoxa  Bugbee:  Hoplitis  producta 
Eurytoma  apiculae  Bugbee:  Ceratina  dallatorreana, 

Ceratina  nanula,  Ceratina  punctigena 
Eurytoma  heriadi  Zerova:  Heriades  crenulatus 
Eun/toma   nodularis   Boheman:   Ceratina   callosa, 

Hylaeus  sp.,  Osmia  adunca,  Osmia  coerulescens, 

Osmia  parvula, 
Eun/toma  sp.:  Osmia  leucomelana 
Eurytoma   stigmi   Ashmead:    Hoplitis   producta, 

Hylaeus  polifolii 

Leucospidae 

Eeucospis  affinis  Say:  Anthidiellum  perplexum, 
Anthidium  emarginatum,  Anthidium  maculo- 
sum,  Ashmeadiella  aridula  astragli,  Ashmeadiella 
meliloti,  Dianthidium  pudicum,  Hoplitis  pro- 
ducta, Megachile  brevis,  Megachile  geutilis, 
Megachile  inermis,  Megachile  mendica,  Mega- 
chile montivaga,  Megachile  pugnata,  Megachile 
relativa,  Osmia  atriventris,  Osmia  californica, 
Osmia  kincaidii,  Osmia  pumila,  Osmia  rostrata, 
Osmia  simillima,  Stelis  sexmaculata. 
Eeucospis  africana  Cameron:  Serapista  denticulata 
Eeucospis  anthidioides  Westwood:  Xylocopa  sub- 

mordax 
Eeucospis  aruina  Walker:  Megachile  rangii 
Eeucospis   bifasciata    Klug:   Anthidiellum   striga- 

tum 
Eeucospis   biguetina  J  urine:   Hoplitis  acuticornis, 
Hoplitis  tridentata 


260 


Journal  of  Hymenoptera  Research 


Leucospis  cayennensis  Westwood:  Centris  analis, 
Centris  bicornuta,  Centris  nitida,  Centris  tarsata, 
Centris  vittata 
Leucospis  dorsigera   ¥.:  Anthidiellum   strigatum, 
Anthidium  diadema,  Anthidium  emarginatum, 
IHeriades  sp.,  Hoplitis  adunca,  Hoplosmia  ligur- 
ica,  Megachile  ericetorum,  Osmia  bicornis,  Osmia 
cornuta,  Osmia  fedtschenkoi,  Osmia  fulviventris, 
Osmia  niveata,  Osmia  parietina,  Osmia  rufa  rufa, 
Osmia  tricornis 
Leucospis  genalis  Boucek:  Diadasina  distincta 
Leucospis  gigas  F.:  Anthophora   bomboides  bom- 
boides,   ICoelioxys   quadridentatus,   Megachile 
hungarica,  Megachile  parietina,  Megachile  pyr- 
enaica,  Megachile  sicula,  Osmia  bicornis,  Osmia 
Icoerulescens,  Osmia  rufa  rufa 
Leucospis  histrio  Maindron:  ICtenoplectra  chaly- 

bea,  Megachile  ustulatum 
Leucospis  hopei  Westwood:  Megachile  rancaguen- 

sis 
Leucospis  intermedia  Illiger:  Megachile  sp.,  Osmia 

emarginata 
Leucospis  japonica  Walker:  Megachile  disjunctifor- 
mis,  Megachile  nipponica,  Megachile  sculpturalis, 
Osmia  excavata,  Osmia  taunts 
Leucospis   klugii   Westwood:   Xylocopa   brasilia- 

norum 
Leucospis  ornata  Westwood:  Lithurgus  capensis, 

Megachile  willowmorensis 
Leucospis  osmaie  Boucek:  Osmia  globicola 
Leucospis  pinna  Grissell  and  Cameron:  Eulaema 

meriana 
Leucospis  poeyi  Guerin-Meneville:  Megachile  poeyi 
Leucospis  reversa  Boucek:  Xylocopa  sp. 
Leucospis  slossonae  Weld:  Anthidiellum  sp. 
Leucospis  sp.:  Microthurge  corumbae 
Leucospis  tricolor  Kirby:   P achy  anthidium  corda- 
tum,  V achy  anthidium  truncatum,  Serapista  den- 
ticulata 
Leucospis  varicollis  Cameron:  Lithurgus  capensis 
Leucospis  xylocopae  Burks:  Xylocopa  nogueirai 
Micrapion  dalyi  Boucek:  Ceratina  sp. 
Micrapion  nasutum  Boucek:  Ceratina  sp. 
Micrapion  richardsi  Boucek:  Ceratina  sp. 
Micrapion  steffani  Boucek:  Ceratina  truucata 
Polistomorpha  conura  Boucek:  Euglossa  sp. 
Polistomorpha  fasciata  Westwood:  Euglossa  ignita, 
Euglossa  sp. 

Mymaridae 

Anagrus  putnamii  Packard:  Megachile  centuncu- 
lari 


Perilampidae 

Aperilampus  varians  Strand:  Halictus  africauus 
Pteromalidae 

Ablaxia  cupraeus  (Provancher):  Megachile  albitar- 

sis,  Megachile  centuncularis 
Cleonymus  amabilis  Cockerell:  Hoplitis  producta, 

Stelis  sexmaculata 
Cleonymus  ceratinae  Kamijo:  Ceratina  japonica 
Cricellius  megachilis  Ashmead:  Megachile  brevis 
Dibrachys  boarmiae  (Walker):  Apis  mellifera 
Dibrachys  cavus  (Walker):  Bombus  sp.,  Megachile 

argent  at  a 
Dibrachys  confusus  (Girault):  Megachile  rotundata 
Dibrachys  maculipennis  Szelenyi:  Megachile  rotun- 
data 
Dibrachys  pelos  Grissell:  Osmia  nigrifrons 
Dibrachys  relativus  Doganlar:  Megachile  relativus 
Dibrachys  sp.:  Megachile  centuncularis,  Megachile 

pugnata 
Epistenia    coeruleata    Westwood:    Ceratina   sp., 

Osnua  sp. 
Habritys  latrus  Wallace:  Ceratina  dupla 
Lariophagus  obtusus  Kamijo:  Megachile  spissula 
Merisus  sp.:  Coelioxys  octodentata,  Megachile  brevis 
Mesopolobus  bruchophagi  (Gahan):  Nomia  melan- 

deri 
Nasonia  vitripennis  (Walker):  Apis  mellifera 
Pachycrepoideus  vindemmiae  Rondani:  Bombus  sp. 
Pteromalus  analis  Ashmead:  Hylaeus  varifrons 
Pteromalus  apum  (Retzius):  Apis  mellifera,  Mega- 
chile centuncularis,  Megachile  pyrenaica,  Mega- 
chile relativa,  Megachile  rotundata,  Megachile 
willughbiella 
Pteromalus     conopidarum     (Boucek):     Bombus 
agrorum,  Bombus  lapidarius,  Megachile  rotun- 
data 
Pteromalus  macronychivorus  Perez:  Megachile  cen- 
tuncularis 
Pteromalus  veneris  Dalla  Torre:  Megachile  rotun- 
data 

Torymidae 

Adoiitomerus  gregalis  (Steffan):  Pseudoanthidium 

lituratum 
Adontomerus  nesterovi  Zerova:   Pseudoanthidium 

lituratum 
Echthrodape  africana    Burks:    Braunsapis    rolini, 

Braunsapis  rufipes,  Braunsapis  simplicipes 
Echthrodape     papuana      Boucek:      Braunsapis 

unicolor 


Volume  16,  Number  2,  2007 


261 


Microdontomerus  anthidii  (Ashmead):    Dianthi- 

dium  pudicum  consimile 
Microdontomerus  apianus  Grissell:  Megachile  mon- 

tivaga 

Microdontomerus  enigma  Grissell:  Hoplitis  bullifa- 
cies 

Microdontomerus  parkeri  Grissell:  Ashmeadiella 
bigeloviae,  Ashmeadiella  cubiceps,  Ashmeadiella 
gillettei,  Ashmeadiella  rufipes,  Hoplitis  bullifa- 
cies,  Hoplitis  palmarum,  Megachile  brevis,  Osmia 
marginata 

Monodontomerus  acrostigmus  Grissell:  Megachile 
sP. 

Monodontomerus  aeneus  (F.):  Anthidium  florenti- 
num,  Anthophora  retusa,  Ceratina  callosa,  Hopli- 
tis adunca,  Megachile  apicalis,  Megachile  cen- 
tuncularis,  Megachile  concinna,  Megachile  par- 
ietina,  Megachile  rotundata,  Megachile  sicula, 
Osmia  brevicornis,  Osmia  coerulescens,  Osmia 
cornigera,  Osmia  cornuta,  Osmia  emarginata, 
Osmia  fulviventris,  Osmia  latreillei,  Osmia  rufa, 
Osmia  nigrifrons,  Osmia  submicans,  Osmia 
tricornis,  Stelis  nasuta 

Monodontomerus  anthidiorum  (Lucas):  Rhodanthi- 
dium  sticticum 

Monodontomerus  argentinus  Brethes:  Eufriesea 
nigrescens,  Megachile  sp. 

Monodontomerus  bakeri  Gahan:  Megachile  pug- 
nata,  Megachile  relativa,  Megachile  rotundata, 
Osmia  coloradensis,  Osmia  texana 

Monodontomerus  brevicrus  Grissell:  Osmia  ribi- 
floris 

Monodontomerus  dementi  Grissell:  Dianthidium 
heterulkei,  Megachile  rotundata 

Monodontomerus  dianthidii  Gahan:  Dianthidium 
sP. 

Monodontomerus  laticornis  Grissell  and  Zerova: 
Anthidium  florentinum,  Anthidium  septemspino- 
sum,  Apis  mellifera,  Megachile  centuncularis, 
Megachile  rotundata 


Monodontomerus  mandibularis  Gahan:  Anthophora 
abrupta,  Anthophora  bomboides  bomboides,  Meli- 
totna  taurea,  Osmia  cordata 

Monodontomerus  mexicanus  Gahan:  Ancyloscelis 
apiformis,  Anthophora  marginata,  Megachile 
peruviana 

Monodontomerus  montivagus  Ashmead:  Anthidium 
collectum,  Anthidium  emarginatum,  Anthidiun 
Imormonum,  Anthophora  abrupta,  Anthophora 
bomboides  bomboides,  Anthophora  bomboides  neo- 
mexicana,  Anthophora  linsleyi,  lAnthophora  occi- 
dentalis,  lAnthophora  vallorum,  Ashmeadiella  cali- 
fornica,  Bombus  morrisoni,  Dianthidium  curvatum 
sayi,  Dianthidium  pudicum  consimile,  Dianthi- 
dium pudicum,  Hoplitis  anthocopoides  (nest), 
Megachile  centuncularis,  Megachile  relativa,  Mega- 
chile rotundata,  IMelissodes  sp.,  Osmia  cordata, 
Osmia  kincaidii,  Osmia  latisulcata,  Osmia  lignaria, 
Osmia  ribifloris,  Osmia  sanrafaelae,  Osmia  texana, 
Stelis  depressa,  Xylocopa  tabaniformis  orpifex 

Monodontomerus  obscurus  Westwood:  Anthophora 
plumipes,  Hoplitis  adunca,  Megachile  argentata, 
Megachile  centuncularis,  Megachile  cephalotes, 
Megachile  flavipes,  Megachile  lanata,  Megachile 
parietina,  Megachile  rotundata,  Megachile  will- 
ughbiclla,  Osmia  cordata,  Osmia  cornifrons, 
Osmia  latreillei,  Osmia  lignaria,  Osmia  ribfloris, 
Osmia  rufa  rufa,  Osmia  rufa  cornigera,  Osmia 
sanrafaelae,  Xylocopa  fenestrata 

Monodontomerus  osmiae  Kamijo:  Osmia  cornifrons, 
Osmia  excavata,  Osmia  taurus 

Monodontomerus  parkeri  Grissell:  Anthophora 
occidentalis 

Monodontomerus  rugulosus  Thomson:  Megachile 
rotundata 

Monodontomerus  tepedinoi  Grissell:  Osmia  lignaria 

Monodontomerus  thorpi  Grissell:  Anthidium  ma- 
culatum 

Monodontomerus  torchioi  Grissell:  Osmia  lignaria, 
Osmia  sanrafaelae 


262 


Journal  of  Hymenoptera  Research 


frenal  line 


upper 
mesepimeron 


lower 
mesepimeron 


metapleuron 


frenum 


transepimeral 

sulcus         epimeron 


metapleuron 


frenum 


malar 
distance 

11 


intermalar 
distance 


submarginal  vein 


marginal 
vein 


postmarginal 
vein 


10 


,     stiqma 
stigmal         y 

parastigma      vein 


Figs.  1-11.  Torymidae.  1-2,  Mesosoma,  side  (arrow  indicates  anterior  margin  of  metapleuron).  3-4,  Metacoxa, 
side.  5-6,  Propodeum,  dorsal.  7-8,  Mesosoma,  dorsal  (arrow  indicates  frenum).  9,  Fore  wing,  dorsal  (Echthrodape 
africana).  10,  Fore  wing,  dorsal,  showing  venation  terminology.  11,  Head,  anterior,  showing  measurements. 


Volume  16,  Number  2,  2007 


263 


14 


15 


16 


occipital 
carina 


hypostomal 
carina 


17 


18 


Figs.  12-28.  Torymidae.  12-13,  Fore  wing  venation,  dorsal.  14-15,  Head,  anterior,  lines  indicate  malar  and 
intermalar  distances.  12,  14  Echthrodape  papuana.  13,  15  Echthrodape  africana.  16-18,  Head,  posterior  (showing 
carinae).  16,  Microdontomerus.  17,  Pseudotorytnus.  18,  Monodontomerus.  19-20,  Antenna,  side.  19,  Monodontomerus 

thorpi.  20,  Monodontomerus  spp.  21-28,  Head.  21-22,  Monodontomerus  osmiae  (from  Kamijo  1963).  23-24, 
Monodontomerus  anthidiorum.  25-26,  Monodontomerus  mexicanus.  27-28,  Monodontomerus  bakeri. 


264 


Journal  of  Hymenoptera  Research 


basal  cell         costal  cell 


admarginal 
area 


postmarginal  vein 


■*■  i*    /*"/*    /■y'gT^v  - 


Figs.  29-42.  Torymidae,  Manodontomerus  spp.  (except  34,  Pseudotorymus).  29-31,  Fore  wing.  29,  M.  aeneus.  30, 
M.  sp.  31,  M.  dementi.  32-33,  Metasomal  tergum  6.  32,  M.  argentinus.  33,  M.  rugulosus.  34-35,  Hind  femur  and 
tibia,  side.  34,  P.  sp.  35,  M.  aeneus.  36-38,  Hind  femur.  36,  M.  thorpi.  37,  M.  argentinus.  38,  M.  rugulosus.  39^0, 
Fore  leg  (left  side  view,  right  ventral  view).  39,  M.  aeneus.  40,  M.  brevierus.  41-42,  Frenum  (apex  of  scutellum).  41, 
M.  acrostigmus.  42,  M.  aeneus. 


Volume  16,  Number  2,  2007 


265 


.  - Ti  i  •> » 


&^scz<'<*s% 


43 


\^> 


51 


52 


\\V^ 


e 


■ 


t 


'.-  ■:'."     I 


transepimeral 
sulcus 


Figs.  43-58.  Torymidae,  Motwdontomerus  (Mo.)  and  Microdontomerus  (Mi.).  43-46,  Fore  wing,  part.  43,  Mo. 
mandibularis.  44,  Mo.  parked.  45,  Mi.  parkeri.  46,  Mi.  enigma.  47-50,  Heads.  47,  Mo.  montivagus.  48,  Mo.  mandibularis. 
49,  Mi.  anthidii.  50,  M/'.  apiamis.  51-52,  M.  acrostigmus  (variation  in  stigma).  53-55,  Mesopleuron.  53,  Mo.  dementi. 
54,  Mo.  parkeri.  55,  Mo.  montivagus.  56-58,  Scape,  male.  56,  Mo.  montivagus.  57,  Mo.  mandibularis.  58,  Mo.  tepedinoi. 


J.  HYM.  RES. 

Vol.  16(2),  2007,  pp.  266-276 

Multivoltinism  and  Usage  of  Multiple  Nest  Substrates  in  a  West  Texas 

Sand  Dune  Population  of  Pseudomasaris  phaceliae  Rohwer 

(Hymenoptera:  Vespidae:  Masarinae) 

John  L.  Neff  and  Allan  W.  Hook 

(JLN)  Central  Texas  Melittological  Institute,  7307  Running  Rope,  Austin,  Texas  78731,  USA; 

email:  jlnatctmi@yahoo.com 
(AWH)  Department  of  Biology,  St.  Edward's  University,  Austin,  Texas  78704-6489,  USA; 

email:  allanh@stedwards.edu 


Abstract. — A  west  Texas  population  of  Pseudomasaris  phaceliae  was  found  to  be  multivoltine  and 
active  from  April  to  September.  Unlike  previous  reports  of  nests  constructed  only  on  stones,  nests 
were  commonly  constructed  on  the  stems  and  infructescences  of  its  host  plant,  Phacelia  integrifolia. 
Emergence  data  indicated  the  primary  sex  ratio  is  strongly  female  biased  (53:14).  Nest  parasitism 
was  rare,  but  predation  was  common.  Data  on  nest  architecture,  nest  construction,  and  foraging 
behavior  are  presented. 


Pseudomasaris  is  the  only  North  Ameri-  phaceliae,  like  most  other  Pseudomasaris 
can  genus  of  the  Masarinae,  a  relatively  species,  constructs  its  nests  on  stones 
small  (300+  spp.)  but  widespread  clade  of  (Parker  1967,  Torchio  1970),  we  found  that 
pollen  and  nectar  provisioning  vespid  at  Monahans,  P.  phaceliae  was  commonly 
wasps  (Carpenter  1982,  2001,  Gess  1996).  attaching  its  nests  to  plant  stems,  and  only 
Torchio  (1970)  reported  in  considerable  rarely  to  stones.  In  addition,  P.  phaceliae, 
detail  on  the  biology  of  Pseudomasaris  like  most  other  temperate  masarine  spe- 
edwardsii  (Cresson)  but  little  is  known  of  cies,  had  previously  been  assumed  to  be 
the  biology  of  the  remaining  14  species,  univoltine  (Parker  1967,  Gess  1996),  but 
Brief  reports  on  nest  structure  and /or  nest  wasps  emerged  in  early  July  from  a  nest 
sites  have  been  published  for  eight  of  these  collected  during  June  suggesting  multi- 
species,  including  Pseudomasaris  phaceliae  voltinism.  This  prompted  a  series  of  visits 
Rohwer,  but  only  floral  records  are  avail-  to  this  site  to  gather  additional  information 
able  for  four  other  species,  and  nothing  at  on  the  nests  and  behavior.  Follow-up  trips 
all  has  been  published  on  the  biology  of  the  were  made  in  July,  August,  and  September 
remaining  two.  2005  and  April,  May,  June,  and  August 

During  a  brief  visit  to  Monahans  Sand-  2006. 

hills  State  Park  in  June  2005,  we  encoun-  Habitat. — Monahans  Sandhills  State  Park 

tered  a  large  population  of  Pseudomasaris  (32.128'N,  103.953  W)  is  located  in  Ward 

phaceliae  visiting  flowers  of  Phacelia  integri-  County,  Texas  on   the  southern  edge  of 

folia   Torr.    (Boraginaceae).    Pseudomasaris  a  large  dune  field  of  quartz-rich  Quaterna- 

phaceliae  is  an  infrequently  collected  but  ry    sands    that    stretch    northward    into 

sometimes  locally  abundant  species  of  the  southeastern   New    Mexico    (Machenberg 

arid  American  Southwest  (Arizona,  New  1984,  Muhs  2001).  While  much  of  the  dune 

Mexico,  west  Texas,  and  adjacent  Mexico)  field    is   partially   stabilized    by   shin   oak 

(Richards  1966,  and  pers.  obs.).  Although  (Quercus  havardii  Rydb.)  and  other  peren- 

previous   reports   had    indicated    that   P.  nials,  large  moving  dunes  are  common  in 


Volume  16,  Number  2,  2007 


267 


Figs.  1-2.     1.  Pseudomasaris  phaceliae  nest  (indicated  by  arrow)  on  Phacelia  integrifolia  stem.  2.  Incomplete  P. 
phaceliae  nest  with  one  completed  cell  and  one  cell  under  construction. 


the  Park.  Soils  in  the  sandhills  consist 
almost  exclusively  of  loose  sands.  Occa- 
sionally, winds  expose  the  underlying 
caliche  layer,  but  generally  there  are  no 
rocks  or  stones  in  the  dunes  beyond  those 
brought  in  for  the  caliche  service  roads. 
Like  most  of  Texas,  summers  are  warm 
with  July  maximum  temperatures  averag- 
ing 35  C.  Rainfall  is  low,  averaging  33.6  cm 
per  year,  with  75%  of  precipitation  occur- 
ring during  a  six-month  May  to  October 
summer/fall  period.  Despite  the  relative 
aridity,  the  water  table  is  quite  shallow  in 
the  sand  hills  due  to  an  underlying 
impermeable  caliche  layer.  Water-loving 
plants  like  Salix  nigra  Marshall  (Salicaceae) 
and  Baccharis  salicina  Torrey  &  A.  Gray 
(Asteraceae)  are  found  in  some  of  the 
deeper  depressions  among  the  dunes 
where  temporary  ponds  may  form  follow- 
ing unusually  heavy  rains  (Machenberg 
1984). 


Nests. — Although  a  few  nests  were 
found  on  miscellaneous  plant  stems  (such 
as  a  sapling  of  Prosopis  glandulosa  Torr. 
(Fabaceae)  or  stems  of  the  erect  herb 
Mentzelia  strictissima  (Wooton  &  Standi.)  J. 
Dark  (Loasaceae),  the  vast  majority  of  the 
nests  we  found  were  on  stems  or  infruc- 
tescences  of  larger  (over  40  cm  high) 
individuals  of  Phacelia  integrifolia  (Fig.  1.). 
Nests  were  located  at  heights  of  20.3- 
71.1  cm  above  the  ground  (n  =  50,  mean  = 
41.0  ±  9.9  cm).  Typically  there  was  only 
one  nest  per  plant  although  a  few  plants 
had  two,  and  one  Phacelia  plant  had  four 
nests,  two  older  nests  from  which  emer- 
gence was  complete,  and  two  newer  nests 
under  construction. 

Like  those  of  other  Pseinloniasaris  species, 
nests  consisted  of  one  or  more  cylindrical 
cells  attached  lengthways  one  to  another 
(Torchio  1970).  Cells  of  nests  on  plants 
were  always  positioned  with  the  long  axis 


268 


Journal  of  Hymenoptera  Research 


oriented  vertically,  regardless  of  the  orien- 
tation of  the  stem  or  infructescence  (Fig.  2). 
Nests  on  plants  averaged  5.4  ±  3.8  cells  per 
nest,  (1-14,  n  =  53),  a  value  that  probably 
underestimates  the  true  number  of  cells 
per  nest  since  some  of  the  nests  may  not 
have  been  complete  when  censused.  Nests 
on  stems  were  initiated  by  laying  down 
a  strip  of  the  sand-soil  mix  along  a  stem  or 
infructescence.  The  female  then  con- 
structed the  hemispherical  inner  end  of 
the  cell  at  the  upper  end  of  the  strip,  and 
completed  the  cell  wall  by  adding  irregular 
strips  or  scale-like  patches  of  the  moist 
sand-soil  mix.  Upon  returning  with  a  sand- 
soil  load,  the  female  inserted  her  head  in 
the  nest  and  curled  her  metasoma  so  its  tip 
was  opposite  her  head.  The  moist  sand-soil 
mix  was  added  to  the  cell  wall  by 
simultaneously  working  the  mix  from 
within  with  her  mandibles  and  tapping  it 
from  the  outside  with  the  flattened,  hirsute 
surface  of  metasomal  sternite  6.  Time  on 
the  nest  depositing  cemented  sand  was 
typically  brief  (0.6-2.9  min). 

The  first  cell  is  a  true  cylinder  as  the 
walls  are  complete,  the  plant  substrate  not 
being  used  as  part  of  the  cell  wall  (Fig.  2). 
Additional  cells  are  attached  to  the  walls  of 
the  initial  cell  so  the  nest  grows  in  an 
asymmetrical  manner  away  from  the  stem 
rather  than  around  it  (Fig.  3).  Cell  walls  are 
rather  thin,  only  0.4-0.5  mm  thick,  but 
quite  strong.  Nests  within  100  meters  of 
the  caliche  road  were  usually  constructed 
with  a  mix  of  relatively  coarse  sand 
particles  (0.1-0.4  mm  in  diameter),  and 
much  finer  (0.02-0.04  mm)  particles,  pre- 
sumably caliche  dust.  Nests  further  from 
the  road  are  constructed  of  sand  alone.  The 
particles  are  bound  together  with  regurgi- 
tated nectar,  and  perhaps,  glandular  prod- 
ucts. As  there  was  no  free  water  in  the 
dunes,  nectar  (or  perhaps  honey-dew)  was 
the  only  likely  source  for  the  bulk  of  the 
liquid  used  to  moisten  the  sand-soil  mix. 
Nest  fragments  placed  in  water  softened 
but  were  still  intact  after  being  immersed 
for   48  hours,   suggesting   substances   be- 


yond just  sugar  may  be  holding  the  walls 
together.  These  moistened  nests  quickly 
sprouted  fungal  hyphae,  the  fungi  pre- 
sumably growing  on  the  nectar  sugars.  In 
contrast,  the  true  mud  nest  walls  and 
partitions  of  nests  constructed  by  Trypargi- 
lum  politum  (Say)  or  Osmia  lignaria  (Say) 
dissolve  almost  instantly  when  placed  in 
water  (pers.  obs.). 

Individual  cells  were  15.2  ±  1.2  mm 
(12.8-17.6,  n  =  19)  mm  long  with  an 
average  diameter  of  4.2  ±  0.2  mm  (3.8- 
4.5,  n  =  28).  The  distal  end  of  the  cell  was 
hemispherical  while  the  cell  opening  was 
simple  and  truncate.  After  provisioning, 
each  cell  was  closed  with  a  cemented  sand 
plug  0.5-0.8  mm  thick  medially  and  1.0- 
1.3  mm  thick  at  the  sides.  Additional  sand 
and  fine  particles  were  added  to  the  nest 
exterior  as  construction  proceeded,  filling 
the  spaces  between  the  cells  (Fig.  9).  In  one 
unfinished  nest  in  which  the  second  cell 
was  only  half  finished,  soil  had  been  added 
along  the  juncture  of  the  two  cells, 
strengthening  their  connection,  although 
additional  soil  had  not  been  added  else- 
where to  the  outer  surface  of  the  first  cell. 
Upon  completion,  the  nest  has  flattened, 
relatively  smooth  walls  concealing  the 
outlines  of  individual  cells.  We  found  no 
indication  of  empty  spaces  between  the 
cells.  Completed  nests  on  stems  usually 
lacked  obvious  ornamentation  but  a  few 
(4  of  50)  had  conical  projections  on  the 
margins  of  the  nest  (Fig.  4). 

Females  were  commonly  observed  col- 
lecting fine  soil  particles  along  the  caliche 
road  (Fig.  6)  and,  less  frequently,  in  the 
dunes.  Females  would  hover  10-15  cm 
above  the  soil  surface  and  repeatedly  drop 
to  collect  sand  or  soil.  We  did  not  obtain 
a  complete  picture  of  soil  collection  but  it 
was  clear  that  during  most  bouts  on  the 
soil  surface,  the  wasps  did  not  add  nectar 
to  the  soil  surface  prior  to  collection. 
Rather,  they  used  their  mandibles  and 
foretarsi  to  add  soil  or  sand  to  a  moist  soil 
bolus  held  behind  the  mandibles  by  their 
modified  labial  palpi.  Regurgitated  nectar 


Volume  16,  Number  2,  2007 


269 


Figs.  3-5.  3.  Completed  Pseudomasaris  phaceliae  nest  showing  asymmetrical  position  on  Phacelia  integrifolia 
stem.  4.  Ornamented  P.  phaceliae  nest  (projection  indicated  by  arrow).  5.  P.  phaceliae  nest  opened  laterally, 
probably  by  birds. 


is  apparently  added  to  this  bolus  while  the 
wasps  are  hovering.  As  in  Pseudomasaris 
edwardsii,  females  constructing  cells  or 
adding  sand  to  the  nest  exterior  typically 
made  long  trips  (20-30  min  duration)  to 
gather  nectar  at  flowers  as  well  as  sand, 
interspersed  between  series  of  3^4  relative- 
ly short  (0.9-3.1  min)  trips.  The  latter  trips 


were  presumably  for  sand  only  since  they 
were  too  brief  to  allow  for  both  sand  and 
nectar  collection. 

While  the  majority  of  nests  were  con- 
structed on  plant  stems,  some  nests  were 
constructed  on  small  stones  brought  in  for 
a  caliche  service  road  through  the  dunes. 
These  were  structurally  similar  to  the  nests 


270 


Journal  of  Hymenoptera  Research 


- 


Figs.  6-10.  6.  Pseudomasaris  phaceliae  female  collecting  sand  (sand  bolus  indicated  by  arrow).  7.  Completed  P. 
phaceliae  nest  on  stone.  8.  Female  extending  walls  of  new  P.  phaceliae  nest.  9.  Female  adding  sand  (indicated  by 
arrow)  to  exterior  of  P.  phaceliae  nest.  10.  Emergence  holes  from  six-  celled  P.  phaceliae  nest. 


constructed  on  stems,  but  had  fewer  cells, 
never  more  than  three,  averaging  only  1.3 
±  0.5  cells  (1-3,  n  =  28).  While  all  the  stem 
nests  were  constructed  during  the  year  in 
which  they  were  discovered,  many  of  the 
rock  nests  were  worn  and /or  damaged 
and  may  have  been  constructed  in  prior 
years.  Unlike  the  situation  in  stem  nests, 
the  first  cells  constructed  on  stones  were 
usually  not  true  cylinders  as  the  stone  was 
used  as  part  of  the  cell  wall.  The  first  cells 
constructed  typically  were  attached  to  the 
stone  along  their  entire  length  (Fig.  8), 
although  in  a  few  cases,  only  the  basal  half 
of  the  cell  was  attached  to  the  substrate, 
with  the  outermost  portion  arching  away. 
Completed  nests  on  stones  often  appeared 
to  be  triangular  in  cross-section,  due  to  the 
smoothing  of  the  sides  and  the  addition  of 
soil,  which  greatly  widened  the  base  of  the 
nest  (Fig.  7).  Nests  on  rocks  occurred  both 
on  the  sides  and  upper  surfaces  without 
any    consistent   orientation.    Completed 


nests  on  stones  were  quite  cryptic  as  the 
outer  nest  covering  matched  the  color  of 
the  pale  stone  on  which  they  were  con- 
structed. Nests  on  Phacelia  plants  were 
more  conspicuous  since  the  pale  nests 
contrasted  with  the  green  stems  or  young 
infructescences,  but  the  nests  were  still 
somewhat  cryptic  since  the  infructes- 
cences, leaves  and /or  leaf  tips  all  com- 
monly turn  brown  with  age. 

Nests  are  provisioned  with  a  dense  mass 
of  regurgitated  pollen-nectar  pellets.  The 
pellets  had  short  projections  and  were 
oriented  so  the  projections,  rather  than 
the  main  mass  of  the  pellets,  contacted  the 
cell  walls.  Pellets  averaged  0.6-0.7  mm 
across  with  projections  from  0.2  to 
0.6  mm  long.  Unlike  the  provisions  of 
Pseudomasaris  edwardsii,  the  innermost  face 
of  the  provision  mass  of  P.  phaceliae  was 
not  smoothed  so  the  individual  pellets 
were  easily  distinguished.  The  outer  face 
of  the  provision  mass  (the  surface  facing 


Volume  16,  Number  2,  2007 


271 


the  cell  closure)  was  smooth  and  convex. 
We  found  several  cells  being  provisioned, 
and  others  with  feeding  larvae  but  re- 
covered only  a  single  cell  with  a  completed, 
intact,  provision  mass.  This  mass  was 
15.2  mm  long,  occupying  nearly  all  the 
cell,  leaving  a  small  open  space  of  0.8  mm 
between  the  mass  and  the  roof  of  the 
hemispherical  inner  end,  and  1.6  mm 
between  the  mass  and  the  cell  closure. 

Host  Plant  and  Foraging. — Pseudomasaris 
phaceliae  is  believed  to  restrict  its  foraging 
to  flowers  of  Pliacelia  spp.  (Torchio  1970). 
At  Monahans  dunes,  individuals  of  P. 
phaceliae  foraged  only  at  flowers  of  Pliacelia 
integrifolia,  the  only  Pliacelia  species  flower- 
ing. Pliacelia  integrifolia  is  a  widespread 
annual  of  the  American  Southwest,  occur- 
ring on  a  variety  of  rocky  or  sandy 
substrates,  particularly  gypsum  or  lime- 
stone (Correll  and  Johnston  1970).  The 
plants  bear  scorpioid  cymes  of  small,  pale 
purple  flowers.  In  west  Texas,  flowering 
occurs  primarily  from  March  through  May 
(Correll  and  Johnston  1970),  although 
herbarium  records  from  the  Plant  Re- 
sources Center  of  the  University  of  Texas, 
Austin,  Texas,  show  flowering  as  late  as 
the  end  of  July  in  the  Monahans  Sandhills 
area,  and  a  non-technical  guide  indicates 
flowering  through  September  (Ajilvsgi 
2003).  Pliacelia  integrifolia  was  common 
and  in  flower  during  our  first  2005  visit 
to  Monahans  dunes  on  16  April,  although 
cold,  wet  weather  prohibited  any  observa- 
tions of  floral  visitors.  When  we  revisited 
Monahans  Dunes  on  18  June  2005,  we 
found  P.  integrifolia  flowering  was  still 
widespread  although  most  abundant  in 
certain  depressions  between  the  dunes. 
When  the  site  was  revisited  a  month  later 
in  July,  most  P.  integrifolia  plants  on  the 
dunes  were  brown  and  dead,  and  flower- 
ing was  restricted  to  the  large  plants 
occurring  in  the  depressions  between  the 
dunes.  Unlike  the  typical,  erect  plants  on 
the  dunes  and  most  other  habitats  (Fig.  1), 
these  plants  were  sprawling  and  multi- 
branched,  almost  shrub-like  in  aspect,  with 


greatly  enlarged  stems.  These  depression 
plants  were  still  strongly  flowering  when 
the  site  was  revisited  in  August.  On  the 
final  2005  visit  on  16-18  September  low 
levels  of  flowering  were  still  occurring 
among  the  large  depression  plants  al- 
though most  plants  were  dead,  and  flower- 
ing was  limited  to  a  few  inflorescences.  On 
21  April  2006,  P.  integrifolia  was  flowering 
and  locally  abundant  on  the  dunes,  but 
surprisingly,  was  absent  in  the  depressions 
where  the  large,  long-flowering  individu- 
als were  found  in  2005.  Flowering  contin- 
ued, with  a  gradual  decline  in  overall 
abundance  through  May  and  June  2006. 
By  August  2006,  only  secondary  inflores- 
cences of  the  few  surviving  individuals 
were  still  flowering. 

Individual  flowers  of  Pliacelia  integrifolia 
begin  opening  around  0830  CDT  (approx. 
2  hrs  after  sunrise),  and  flowers  continue 
opening  through  the  day.  Individual  flow- 
ers last  approximately  two  days.  Anthers 
dehisce  fully  shortly  after  flower  opening, 
and  are  usually  stripped  of  pollen  within 
two  hours  of  opening.  Some  larger  bees, 
such  as  species  of  Habropoda,  Osmia,  or 
Martinapis  began  foraging  on  P.  integrifolia 
at  or  before  0800  early  in  the  season,  but 
Pseudomasaris  phaceliae  was  always  a  late 
starter.  Females  were  rarely  observed 
before  0830  CDT  and  they  were  generally 
not  active  until  after  0930  CDT  with  air 
temperatures  above  30  C.  Foraging  contin- 
ued until  sundown,  although  females  were 
sometimes  observed  resting  on  stems  or 
branches  during  periods  of  peak  heat  in 
late  afternoon  (air  temperatures  above 
39  C). 

Foraging  rates  recorded  on  13  July  2005 
were  quite  rapid,  perhaps  a  reflection  of 
the  high  temperatures  (33-35  C)  and  re- 
source depletion  due  to  the  high  number  of 
wasps  and  bees  concentrated  on  the  rela- 
tively few  Pliacelia  plants  still  flowering. 
Wasps  visited  an  average  of  24.5  ±  5.3 
(16.4-31.2,  n  =  20)  Pliacelia  flowers  per 
minute.  Nectar  visits  were  quite  quick, 
averaging  only  0.8  ±  0.3  (0.3-2.0,  n  -  40) 


272 


Journal  of  Hymenoptera  Research 


Figs.  11-12.     11.  Pseudomasaris  phaceliae  female  collecting  pollen  of  Phacelia  integrifolia.  12.  P.  phaceliae  female 
nectaring  at  flower  of  P.  integrifolia. 


seconds  per  flower  while  pollen-collecting 
visits  were  longer  at  4.1  ±  3.0  (1.1-11.2,  n  = 
20)  seconds.  During  a  foraging  bout,  the 
proportion  of  flowers  worked  for  pollen 
was  low,  averaging  only  13.7  ±  14.6%  (0- 
41.7,  n  =  20).  This  is  presumably  a  re- 
flection both  of  the  low  availability  of  fresh 
flowers  with  available  pollen,  and  the  fact 
that  some  foragers  were  constructing  their 
nests  and  not  foraging  for  pollen  at  the 
time.  If  we  exclude  foraging  series  where 
no  flowers  were  visited  for  pollen,  the 
proportion  of  flowers  visited  for  pollen 
rises  only  slightly  to  19.6  ±  13.7%  (3.4-41.7, 
n  =  14). 

A  nectar  foraging  bout  involved  landing 
on  medial  portions  of  the  exserted  staminal 
filaments  and  rapidly  inserting  the  re- 
markably extensible  proboscis  (~  4.5  mm 
long  when  fully  extended)  to  reach  the 
nectary  at  the  base  of  corolla  tube  (Fig.  12). 
The  initial  approach  to  a  flower  presum- 
ably involved  some  assessment  of  pollen 
availability,  since  foraging  behavior  chan- 
ged when  a  pollen  forager  encountered 
a  flower  with  obvious  available  pollen.  A 
pollen  forager  would  grasp  the  staminal 
filament  near  the  anther  with  her  hind  and 
mid-tarsi  while  hovering  (Fig.  11).  She 
would    then   grasp    an   anther   with   her 


mandibles  and  extract  pollen  by  scraping 
pollen  to  her  mouthparts  with  the  tarsal 
brushes  of  her  forelegs.  Depending  on 
pollen  availability,  a  wasp  might  work 
several  anthers  on  a  flower  before  moving 
down  the  filaments  to  insert  her  mouth- 
parts  to  gather  nectar. 

A  female  observed  on  17  June  2006  took 
8.40  hrs  and  15  foraging  trips  to  provision 
a  cell.  Pollen  foraging  appeared  to  com- 
mence immediately  after  oviposition.  Be- 
cause of  poor  light  conditions,  we  were  not 
able  to  determine  if  the  female  deposited 
an  initial  pollen  and  nectar  load  immedi- 
ately after  ovipositing  as  has  been  reported 
for  Pseudomasaris  edwardsii  (Torchio  1970). 
Pollen  trips  averaged  31.53±7.10  min 
(19.32-41.72,  n  =  13)  and  deposition  time 
in  the  nest  between  trips  averaged 
2.06±0.48  min  (1.50-3.52,  n  =  14).  As  noted 
for  P.  edwardsii  (Torchio  1970),  the  female 
rotated  within  the  cell  during  the  later 
phases  of  pollen  deposition  as  she  de- 
posited pollen  pellets. 

Development  and  Voltinism. — Only  frag- 
mentary data  were  obtained  on  develop- 
ment. The  only  intact  egg  recovered  was 
smooth,  slightly  asymmetrical,  and  3.7  mm 
long  with  a  medial  width  of  1.0  mm.  The 
posterior   end    of   the   egg   was   attached 


VI 


Ml 


tfl 


Volume  16,  Number  2,  2007 


273 


Table  1.     Collection  dates  for  nests  of  Pseudomasaris  phaceliae  from  Monahans  Sandhills  with  emergence  dates 
ind  sex  ratio  of  the  wasps. 


Nest 

Collection  date 

Emergence  date 

Males 

Females 

H-l 

16-vi-2005 

by  12-vii-2005 

1 

14 

M-l 

18-vii-2005 

19-vii-2005 

0 

2 

\J-2 

18-vii-2005 

23-vii-2005 

0 

1 

SJ-3 

18-vii-2005 

3-iv-2006 

0 

1 

vJ-4 

15-viii-2005 

28-viii-2005 

1 

3 

M-5 

15-viii-2005 

l-ix-2005 

0 

1 

sr-6 

18-vii-2005 

19-iv-2006 

0 

2 

)5-06 

18-vii-2005 

approx.  3-iv-2006 

0 

1 

)5-07 

18-vii-2005 

approx.  3-iv-2006 

1 

1 

15-02 

18-vii-2005 

18-iv-2006 

2 

0 

)6-01 

23-iv-2006 

26  to  29  iv-2006 

1 

5 

)6-02 

19-V-2006 

1  to  7  vi-2006 

1 

8 

16-03 

18-vi-2006 

27-vi  to  11 -vii-2006 

1 

7 

16-04 

18-vi-2006 

1  to7  vii-2006 

4 

3 

16-05 

18-vi-2006 

5  toll  vii-2006 

1 

3 

16-06 

18-vi-2006 

12-vii-2006 

1 

1 

fotal 

14 

53 

perpendicularly  to  the  inner  surface  of  the 
:ell  wall,  just  below  the  hemispherical, 
nner  end  of  the  cell.  The  ventral  surface  of 
he  egg,  which  paralleled  the  flat  inner 
;urface  of  the  provision  mass,  was  some- 
vhat  flattened,  but  the  dorsal  surface  of  the 
?gg  arched  slightly  into  the  curved  space  of 
he  hemispherical  inner  cell  cap.  In  two 
nstances  where  late  instar  larval  feeding 
vas  noted,  the  larva  fed  along  the  side  of 
he  provision,  eating  its  way  down  the 
provision  mass  toward  its  distal  end. 
Sometime  after  completing  feeding,  the 
arva  spun  a  thin,  translucent  cocoon  that 
idhered  tightly  to  the  cell  wall.  Defecation 
>ccurred  after  the  completion  of  the  co- 
:oon.  Feces  were  typically  deposited  as  an 
rregular  ring  of  smooth,  flattened,  semi- 
.pherical  pellets  (0.8-1.0  mm  wide,  and 
1.5-0.6  mm  tall),  around  the  inner  end  of 
he  cell,  although  some  of  these  are 
ometimes  pressed  into  a  flattened  cake 
vith  unrecognizable  individual  pellets. 
\Iest  dissections  indicated  wasps  overwin- 
er  as  prepupae.  Adults  emerge  by  chew- 
ng  through  the  nest  plug  (Fig.  10). 

The  total  number  of  generations  per  year 
)f  Pseudomasaris  phaceliae  at  Monahans  is 
inknown  but  at  a  minimum  it  is  two  and 


perhaps  as  many  as  four.  Pseudomasaris 
phaceliae  was  active  at  Monahans  for  at 
least  93  days  (18  June  to  18  September)  in 
2005  and  116  days  (26  April  to  19  August) 
in  2006.  Judging  from  the  extensive  wing 
wear  of  females  collected  in  June  of  2005, 
the  phenology  of  PJiacelia  iutegrifolia,  and 
the  emergence  times  of  P.  phaceliae  in  the 
lab  (Table  1.),  flight  at  Monahans  during 
2005  probably  began  in  early  April  sug- 
gesting a  flight  period  in  excess  of  156  days 
in  2005. 

Emergence  patterns  of  wasps  from  nests 
collected  in  2005  and  2006  are  indicated  in 
Table  1.  All  nests  collected  before  July  had 
their  inhabitants  emerge  that  same  year, 
but  for  nests  collected  in  July  or  later,  some 
emerged  the  same  year  but  others  went 
into  larval  diapause  and  emerged  the 
following  April.  The  instances  of  late  2005 
(July  to  September)  emergence  were  from 
nests  taken  from  infructescenses  on  rela- 
tively fresh  green  plants,  and  it  is  clear  that 
they  had  been  provisioned  in  2005.  Phacelia 
iutegrifolia  stems  usuallv  break  down  com- 

O      J  J 

pletely  over  the  winter  and  the  glandular 
epidermis,  to  which  the  nests  are  frequent- 
ly attached,  falls  away  soon  after  the  plant 
dies.    It   is   probable   that   over-wintering 


274  Journal  of  Hymenoptera  Research 

nests  fall  from  the  disintegrating  plants  was    not    confirmed.    Additionally,    two 

and  spend  the  winter  in  the  sand.  newly  provisioned  cells  were  encountered 

Mating  and  Sex  Ratio.— Males  of  Pscudo-  with  small  lateral  slits,  possibly  the  results 

masaris  phaceliae  were  observed  to  forage  of  bird  probes.  One  of  these  probed  cells 

for  nectar  at  plants  of  Phacelia  integrifolia  contained  a  larva  and  partially  consumed 

and  patrol  P.  integrifolia  inflorescences,  but  pollen  mass   while   the  other  was  being 

mating  was  not  observed.  Newly  emerged  raided  and  emptied  by  ants, 

males  confined  with  newly  emerged  fe-  We  found  no  indication  of  empty  (closed 

male  nest  mates  attempted  to  mount  their  but  unprovisioned)  cells  in  Pseudomasaris 

sisters,  although  it  was  not  determined  if  phaceliae  nests  at  Monahans.  Empty  cells 

mating  was  successful.  Females  were  much  have   been    invoked    as   an   anti-parasite 

more  common  than  males  at  flowers  at  all  defense  strategy  for  Pseudomasaris  vespoides 

times  during  our  visits,  and  the  sex  ratio  of  (Cresson),  in  which  roughly  30%  of  the  nest 

wasps  emerging  from  nests  was  heavily  cells  are  empty  (Tepedino  et  al.  1979). 
female  biased  3.8:1  (53  females/ 14  males, 

Table  1).    In   all   cases   where   emergence  DISCUSSION 

order   was   determined,    males   emerged  Our  observations  of  Pseudomasaris  phace- 

from  the  first  provisioned  cell(s)  of  a  nest.  Hae  at  Monahans  suggest  this  species  has 

Nest  Associates  and  Predators. — Nest  par-  a  broader  behavioral  range  than  previously 
asitization  was  uncommon.  One  female  observed  in  Pseudomasaris.  The  most  obvi- 
Chrysurissa  densa  (Cresson)  (Chrysididae)  ous  differences  from  previous  reports  are 
emerged  on  18  July  2005  from  a  Pseudoma-  the  incidence  of  multivoltinism  and  use  of 
saris  phaceliae  nest  and  a  second  female  plant  stems  for  nest  placement.  Multivoltin- 
emerged  from  a  different  nest  on  13  ism  has  not  previously  been  reported  in 
September  2005.  Chrysurissa  densa  appar-  Pseudomasaris  and  appears  to  be  rare  in  the 
ently  is  a  specialist  on  Pseudomasaris,  since  Masarinae,  at  least  among  temperate  zone 
besides  P.  phaceliae,  its  only  known  hosts  species  (Gess  1996).  Presumably,  this  is  true 
are  four  other  Pseudomasaris  species.  In  because  these  wasps  are  typically  oligolectic 
addition,  its  range  mirrors  that  of  Pseudo-  (or  at  least  have  a  narrow  range  for  floral 
masaris  (Bohart  and  Kimsey  1982).  The  only  hosts)  and  the  flowering  periods  of  their 
other  nest  parasites  where  10  females  and  2  floral  hosts  usually  are  quite  temporally 
males  of  an  unidentified  Monodontomerus  restricted.  Multivoltinism  is  possible  at 
sp.  (Torymidae)  which  emerged  from  a  cell  Monahans  because  of  the  unusual  extended 
of  another  nest  of  P.  phaceliae.  As  parasite  flowering  of  its  floral  host  in  this  distinctive 
emergence  occurred  only  two  days  after  habitat,  allowing  a  flight  season  that  can 
collection  of  the  nest,  it  was  clearly  the  extend  from  April  into  September  in  favor- 
result  of  field,  rather  than  laboratory,  able  years.  However,  it  is  possible  that 
infestation.  multivoltinism  occurs  in  other  non-dune 

Indications  of  nest  predation  were  occa-  populations  of  P.  phaceliae  associated  with  P. 

sionally   encountered.   Several   nests   ob-  integrifolia.  We  have  collected  P.  phaceliae  on 

served  in  2005  and  3  of  21  nests  measured  this  species  at  other  west  Texas  sites  in  June, 

in  June  2006  had  all  their  cells  opened  Wing   wear  indicated   these   wasps  were 

laterally  (Fig.  5).  The  cells  in  these  nests  relatively  newly  emerged  even  though  the 

contained  empty  cocoons  but  had  intact  few  flowering  P.  integrifolia  plants  at  these 

cell  caps.  Birds  are  the  most  likely  pre-  sites  were  in  very  poor  condition  and  most 

dators  although  we  cannot  rule  out  small  plants  in  the  populations  were  dead   or 

mammals.    A    small,    unidentified    wood-  fruiting. 

pecker  was  seen  perching  and  searching  As   we   encountered   neither   flowering 

on  Phacelia  stems,  but  actual  nest  predation  Phacelia  nor  Pseudomasaris  phaceliae  during 


Volume  16,  Number  2,  2007 


275 


July,  August,  or  September  visits  to  Mon- 
ahans  prior  to  2005,  multivoltinism  may  be 
a  facultative  phenomenon  for  P.  phaceliae. 
The  wasps  could  use  environmental  cues 
such  as  temperature  and  humidity,  which 
may  predict  extended  Phacelia  bloom,  to 
"determine"  whether  to  pupate  and 
emerge  immediately  or  proceed  to  dia- 
pause. Increasing  humidity,  a  predictor  of 
flowering  in  desert  plants,  has  experimen- 
tally been  shown  to  be  an  important  cue  in 
breaking  diapause  in  Macrotera  portalis 
Timberlake  (Andrenidae),  a  desert  bee 
with  extended  diapause  (Danforth  1999), 
while  temperature  (above  or  below  29°C) 
determines  whether  Nomia  melanderi  Cock- 
erell  (Halictidae)  pupates  and  emerges 
immediately  or  proceeds  to  diapause  (Ste- 
phen 1965).  Even  in  years  with  extended 
Phacelia  flowering,  flowering  declines 
greatly  late  in  the  year.  The  observation 
that  some  larvae  in  late  provisioned  nests 
(July  or  later)  pupate  and  emerge  immedi- 
ately while  others  diapause  until  the 
following  spring  suggest  a  bet-hedging 
strategy  predicated  on  the  decreasing 
chances  of  encountering  adequate  floral 
resources  late  in  the  year. 

Published  reports  have  indicated  that 
surfaces  of  rocks  are  the  preferred  nest 
substrates  utilized  by  six  Pseudomasaris 
species  (Hicks  1929,  Hungerford  1937, 
Parker  1967).  In  Torchio's  1970  glasshouse 
study,  Pseudomasaris  edwardsii  females  con- 
structed nests  on  a  variety  of  substrates, 
but  not  plant  stems.  However,  the  use  of 
bamboo  stakes  as  a  nest  substrate  in 
Torchio's  study  suggests  they  may  occa- 
sionally use  twigs  or  plant  stems  under 
natural  conditions.  The  only  report  on 
Pseudomasaris  texana  (Cresson)  indicates  it 
constructs  its  nests  on  twigs  (Bequaert 
1940).  Pseudomasaris  vespoides  has  repeat- 
edly been  reported  constructing  its  nests 
on  twigs  or  plant  stems  (Cockerell  1913, 
Davidson  1913,  Hicks  1929,  Bequaert  1940, 
Torchio  1970),  although  it  also  is  known  to 
use  rocks  (Hicks  1927).  Pseudomasaris  mar- 
ginalis   (Cresson)    was   found    to   nest   in 


beetle  borings  in  logs  at  a  high  altitude 
site  in  Colorado  (Dorr  and  Neff  1982) 
although  this  report  has  been  questioned 
(Gess  1996).  In  the  only  previous  report  on 
the  biology  of  P.  phaceliae,  rock  surfaces 
were  the  only  reported  nest  substrate 
(Parker  1967). 

Since  rocks  are  rarely  encountered  in  the 
Monahans  Sandhills,  it  is  not  surprising 
that  most  of  the  Pseudomasaris  phaceliae 
nests  we  encountered  were  attached  to 
plants.  The  relative  advantages  of  rock 
surfaces  and  plant  stems  as  nest  substrates 
are  unclear.  Rock  surfaces  are  obviously 
more  permanent  and  stable  than  herba- 
ceous plant  stems,  but  the  importance  of 
this  difference  is  not  clear  for  wasps  like 
these  that  do  not  reuse  their  nests.  The 
issue  of  difference  of  substrate  permanence 
could  easily  be  erased  by  using  the  stems 
of  perennials,  although  P.  phaceliae  does  not 
seem  to  regularly  do  this  at  Monahans.  If 
heat  stress  is  a  problem,  a  strong  possibility 
in  a  habitat  like  Monahans  Sandhills  where 
soil  surface  temperatures  regularly  exceed 
40°  during  the  late  spring  and  summer, 
constructing  nests  well  above  the  soil 
surface  on  plant  stems  might  be  advanta- 
geous relative  to  building  nests  on  low, 
exposed  rock  surfaces. 

The  nest  biology  of  most  Pseudomasaris 
species  is  poorly  known,  often  based  on 
only  a  single  population,  and,  in  some 
cases,  a  single  nest.  When  the  biologies  of 
more  populations  of  more  species  of 
Pseudomasaris  are  known,  it  will  be  in- 
teresting to  see  if  other  Pseudomasaris 
species  are  similarly  flexible  in  their  pat- 
terns of  voltinism  and /or  nest  substrate 


usage. 


ACKNOWLEDGEMENTS 


We  thank  the  Texas  Parks  and  Wildlife  Department 
(Scientific  Study  Permit  27-05)  and  the  staff  at 
Monahans  Sandhills  State  Park  for  the  opportunity 
to  study  the  Monahans  Pseudomasaris  population. 
Beryl  B.  Simpson  (The  University  of  Texas)  improved 
early  drafts  of  the  manuscript.  Sarah  Gess  and  an 
anonymous  reviewer  made  many  useful  comments  on 
the  submitted  manuscript. 


276 


Journal  of  Hymenoptera  Research 


LITERATURE  CITED 

Ajilvsgi,  G.  2003.  Wild/lowers  of  Texas,  revised  edition. 
Shearer  Publishing,  Fredericksburg,  Texas,  xix  + 
524  pp. 

Bequaert,  J.  1940.  Notes  on  the  distribution  of  Pseudo- 
masaris  and  on  the  foodplants  of  the  Masaridinae 
and  Gayellinae  (Hym.,  Vespidae).  Bulletin  of  the 
Brooklyn  Entomological  Society  35:  37-45. 

Bohart,  R.  M.  and  L.  S.  Kimsey.  1982.  Chrysididae  in 
America  North  of  Mexico.  Memoirs  of  the  American 
Entomological  Institute  33:  1-266. 

Carpenter,  J.  M.  1982.  The  phylogenetic  relationships 
and  natural  classification  of  the  Vespoidea  (Hy- 
menoptera). Systematic  Entomology  7:  11-38. 

.  2001.  Checklist  of  species  of  the  subfamily 

Masarinae  (Hymenoptera:  Vespidae).  American 
Museum  Novitates  3325:  1-40. 

Cockerell,  T.  D.  A.  1913.  Pseudomasaris  bred  in 
California.  Proceedings  of  the  Entomological  Society 
of  Washington  15:  107. 

Correll,  D.  S.  and  M.  C.  Johnston.  1970.  Manual  of  the 
Vascular  Plants  of  Texas.  Texas  Research  Founda- 
tion, Renner,  Texas,  xv  +  1881  pp. 

Danforth,  B.  N.  1999.  Emergence  dynamics  and  bet 
hedging  in  a  desert  bee,  Perdita  portalis.  Proceed- 
ings of  the  Royal  Society  of  London,  B  266: 
1985-1994. 

Davidson,  A.  1913.  Masaria  vespoides.  Bulletin  of  the 
Southern  California  Academy  of  Science  12:  17-18. 

Dorr,  L.  J.  and  J.  L.  Neff.  1982.  Pseudomasaris  marginalis 
nesting  in  logs  in  Colorado  (Hymenoptera: 
Masaridae).  Pan-Pacific  Entomologist  58:  124-128. 

Gess,  S.  K.  1996.  The  Pollen  Wasps.  Ecology  and  Natural 
History  of  the  Masarinae.  Harvard  University 
Press,  Cambridge,  Massachusetts,  x  +  340  pp. 

Hicks,  C.  H.  1927.  Pseudomasaris  vespoides  (Cresson), 
a  pollen  provisioning  wasp.  Canadian  Entomologist 
59:  75-79. 


.  1929.  Pseudomasaris  edwardsii  Cresson,  another 

pollen  provisioning  wasp,  with  further  notes  on 
P.  vespoides  (Cresson).  Canadian  Entomologist  61: 
122-125. 

Hungerford,  H.  B.  1937.  Pseudomasaris  occidentalis 
(Cresson)  in  Kansas  (Hymenoptera  -  Vespidae). 
Journal  of  the  Kansas  Entomological  Society  10: 
133-134. 

Machenberg,  M.  D.  1984.  Geology  of  Monahans  Sand- 
hills State  Park,  Texas.  Bureau  of  Economic 
Geology,  The  University  of  Texas  at  Austin. 
39  pp.  ' 

Muhs,  D.  R.  and  V.  T.  Holliday.  2001.  Origin  of  late 
Quaternary  dune  fields  on  the  Southern  High 
Plains  of  Texas  and  New  Mexico.  Geological 
Society  of  America  Bulletin  113:  75-87. 

Parker,  F.  D.  1967.  Notes  on  the  nests  of  three  species 
of  Pseudomasaris  Ashmead  (Hymenoptera:  Masar- 
idae). Pan-Pacific  Entomologist  43:  213-14. 

Richards,  O.  W.  1966.  New  records  of  Pseudomasaris 
Ashmead  (Hymenoptera:  Vespoidea,  Masaridae), 
with  notes  on  P.  phaceliae  Rohwer  and  P.  cazieri  R. 
M.  Bohart.  Proceedings  of  the  Royal  entomological 
Society,  London  (B)  35:  47-55. 

Stephen,  W.  P.  1965.  Temperature  effects  on  the 
development  and  multiple  generations  in  the 
alkali  bee,  Nomia  melanderi  Cockerell.  Entomolo- 
gica  Experimentalis  et  Applicata  8:  228-240. 

Tepedino,  V.  J.,  L.  L.  McDonald,  and  R.  Rothwell. 
1979.  Defense  against  parasitization  in  mud- 
nesting  Hymenoptera:  Can  empty  cells  increase 
net  reproductive  output.  Behavioral  Ecology  and 
Sociobiology  6:  99-104. 

Torchio,  P.  F.  1970.  The  ethology  of  the  wasp, 
Pseudomasaris  edwardsii  (Cresson)  and  a  descrip- 
tion of  its  immature  forms  (Hymenoptera:  Ves- 
poidea, Masaridae).  Los  Angeles  County  Museum 
Contributions  in  Science  202:  1-32. 


IH 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  277-280 

Mydrosoma  micheneri  Packer,  new  species,  a  New  Diphaglossine  Bee 

from  Brazil  (Hymenoptera:  Colletidae) 

Laurence  Packer 

Department  of  Biology,  York  University,  4700  Keele  St.,  Toronto,  Ontario,  M3J  1P3,  CANADA; 

email:  bugsrus@yorku.ca 


Abstract. — Mydrosoma  micheneri  Packer,  new  species,  is  described  and  illustrated.  The  sole 
known  specimen,  a  female,  is  from  the  Mato  Grosso  of  Brasil  and  was  collected  almost  40  years  ago. 
It  is  distinctive  in  having  a  longer  head  and  malar  space  than  other  members  of  its  tribe. 


The  purpose  of  this  paper  is  to  describe 
a  somewhat  unusual  species  in  the  genus 
Mydrosoma.  It  differs  from  other  species  by 
the  comparatively  elongate  malar  space  and 
clypeus.  The  genus  Mydrosoma  is  one  of 
three  genera  in  the  tribe  Dissoglottini,  the 
others  being  Mydrosomella,  with  two  species 
(Graf  and  Urban  2001),  and  the  monotypic 
Ptiloglossidia  (Michener  2007).  Mydrosoma 
occurs  from  southern  Brazil  to  Mexico  and 
none  of  its  nine  species  have  been  collected 
frequently.  The  only  biological  data  suggest 
that  these  may  be  late  afternoon  flying  bees, 
with  short  activity  periods;  this  could  help 
explain  their  apparent  rarity. 

In  the  description  below,  standard  ter- 
minology for  bee  morphology  is  employed, 
following  Michener  (1986,  2007).  Puncture 
density  is  indicated  by  the  relative  dis- 
tances between  punctures  in  terms  of 
interspace  (i)  to  puncture  diameter  (d) 
ratios  (e.g.  i  =  2d).  Flagellomeres  are 
numbered  1-10,  and  metasomal  terga  and 
sterna  indicated  by  T  and  S,  respectively. 
Hair  length  is  indicated  relative  to  the 
diameter  of  the  median  ocellus  -  MOD. 

Mydrosoma  micheneri  Packer  new  species 
Figs  1-2 

Diagnosis. — The  new  species  has  the 
standard  combination  of  characteristics  of 
the  tribe  Dissoglottini:  pre-episternal 
groove    absent    and    notaulus    weak    or 


absent.  It  is  clearly  a  member  of  the  genus 
Mydrosoma  as  indicated  by  the  presence  of 
arolia,  second  and  third  submarginal  cells 
subequal  in  area  and  basitibial  plate  in- 
complete. The  new  species  is  readily 
separated  from  other  Mydrosoma  by  the 
comparatively  elongate  head,  with  clypeus 
only  1.5  times  as  wide  as  long  and  malar 
space  almost  as  long  as  basal  depth  of 
mandible  (Fig.  1). 

Description. — Female.  Body  length  14  mm, 
forewing  length  9  mm,  head  width 
3.05  mm,  intertegular  span  2.8  mm. 

Colouration:  Black  with  lower  face, 
antenna,  legs  and  metasoma  dark  brown; 
following  parts  orange:  anterior  surface  of 
flagellum  (except  F2  red-brown),  entire 
apical  flagellomere,  tegula,  fore  tibia  and 
fore  tarsus,  wing  veins;  wing  membrane 
pale  amber;  metasomal  terga  with  metallic 
reflections;  apical  impressed  areas  straw. 

Pubescence:  Hairs  plumose  with  nu- 
merous short  branches.  Bright  fuscous  on 
dorsal  and  lateral  surfaces  of  mesosoma, 
pale  yellowish  on  face,  ventral  surface  of 
mesosoma,  legs  and  metasoma.  Outer 
surface  of  hind  tibia  with  brown  hairs. 
Prepygidial  fimbria  dark  brown,  hairs  on 
disks  of  T2-T5  blackish.  Hairs  on  face 
short,  1.5MOD;  slightly  longer  on  vertex, 
genal  area  and  mesoscutum,  2MOD;  longer 
on  mesopleuron,  scutellum  and  metano- 
tum    <3MOD;    longest    hairs   on    lateral 


278 


Journal  of  Hymenoptera  Research 


Fig.  1.     Lateral  habitus  of  Mydrosoma  micheneri,  Packer,  n.  sp. 


surface  of  propodeum  and  hind  femoral 
scopa,  4MOD.  Hind  tibia  with  shorter  hairs 
<2MOD  except  longer  on  ventral  surface 
<3MOD.  Tl  with  simple  erect  hairs 
2MOD,  longer  and  plumose  laterally, 
3MOD.  Apical  bands  of  appressed  hairs 
on  T2-T4  <2MOD.  SI  with  short  erect 
hairs,  1.5MOD;  S2-S5  with  posteroven- 
trally  directed  long  subapical  hair  bands, 
hairs  with  numerous  branches  on  anterior 
of  rhachis  only,  longest  on  S3,  3.5MOD. 

Surface  sculpture:  Microsculpture  weak, 
surface  shiny,  except  somewhat  dull  on 
lower  face.  Punctures  on  face  below  anten- 
nae distinct  and  moderately  dense  i  ~  d; 
frons  with  punctures  finer  and  shallower 
but  equally  dense;  narrow  transverse  im- 
punctate  band  between  antennal  bases;  area 
between  lateral  ocellus  and  compound  eye 
and  vertex  immediately  behind  ocelli  al- 
most impunctate.  Mesoscutum  with  shal- 
low, moderately  dense  punctures,  i   ^  d; 


scutellum  with  deeper,  more  distinct  and 
irregularly  spaced  punctures,  i  =  l-3d; 
metanotum  with  punctures  almost  crowded 
laterally,  i  >  d  medially;  dorsal  area  of 
propodeum  impunctate,  lateral  surface 
densely  punctate  above  and  along  ventral 
margin,  i  <  d,  sparsely  punctate  below; 
mesopleuron  with  dense,  somewhat  effaced 
punctures,  almost  crowded  dorsally,  i  < 
1.5MOD  below;  Tl  with  punctures  sparse 
on  disk,  i  =  2-4d,  denser  towards  sub- 
marginal  zone  i  =  l-2d;  punctures  increas- 
ingly dense  on  more  posterior  terga,  i  ^  d  on 
T5.  Apical  impressed  areas  with  tiny  dense 
punctures  apically. 

Structure:  Head  as  wide  as  long.  La- 
bium flat  with  weak  transverse  basal  ridge. 
Mandible  three  times  as  long  as  basal 
depth,  subapical  tooth  short.  Galeal  comb 
with  approximately  28  teeth.  Clypeus  1.5 
times  as  wide  as  long,  apical  rim  slightly 
upturned.  Supraclypeal  area  1.25  times  as 


Volume  16,  Number  2,  2007 


279 


Fig.  2.     Frontal  view  of  head  of  M.  micheneri  Packer  n.  sp.  to  show  elongate  mandible,  clypeus  and  malar  area. 


long  as  apical  breadth.  Interantennal  dis- 
tance 0.75 X  antennocular  distance.  Malar 
space  long,  —0.87X  as  long  as  basal  depth 
of  mandible.  Genal  length  subequal  to 
width  of  compound  eye;  longitudinal  axis 
of  compound  eye  just  posterior  to  mid- 
depth  of  mandible;  almost  at  right  angle  to 
axis  of  mandible  base.  Facial  fovea  in- 
distinct, marked  by  weak  ridge  at  inner 
margin  of  upper  paraocular  area  approxi- 
mately 1.5MOD  in  length.  Ocellocular 
distance  subequal  to  interocellar  distance 
and  less  than  twice  MOD.  Vertex  behind 
lateral  ocellus  subequal  in  length  to  MOD, 
weakly  depressed.  Fl  equal  to  combined 
lengths  of  F2  and  F3;  F2-F9  somewhat 
broader  than  long,  F10  almost  twice  as  long 
as  wide.  Notaulus  not  evident.  Scutellum 
weakly  depressed  medially.  Dorsal  surface 
of  propodeum  convex,  mostly  declivous. 
Basal  vein  thickened  for  apical  half.  First 
recurrent  vein  enters  second  submarginal 


cell  two  vein  widths  from  lr-m.  Posterior 
margin  of  second  submarginal  cell  10% 
longer  than  that  of  third  submarginal  cell. 
Basitibial  plate  indicated  by  posterior 
carina  2.5MOD  in  length.  Hind  basitarsus 
twice  as  long  as  greatest  depth,  dorsal 
margin  strongly  and  ventral  margin  weak- 
ly convex.  Jugal  lobe  slightly  less  than  half 
as  long  as  vannal  lobe. 

Male. — Unknown. 

Etymology. — It  is  a  pleasure  to  name  this 
species  after  Charles  Michener  in  recogni- 
tion of  his  stellar  achievements  in  melittol- 

°gy- 

Material  Studied. — Holotype  female: 
BRAZIL:  Mato  Grosso  12  50'S  51  47'VV, 
2.iv.l968.  O.W.  Richards.  A  second  label 
states:  R.S.  &  R.G.S.  Expedition  B.M.  1968- 
260.  A  third  label  states  "Gallery  forest".  A 
fourth,  handwritten,  label  states  "Paracol- 
letini,  n.  gen.  N.  sp.!"  [Though  the  species 
is  clearly  a  member  of  the  Diphaglossinae 


280 


Journal  of  Hymenoptera  Research 


as  indicated  by  the  tiny  stigma  and 
strongly  bifid  glossa.]  The  specimen  be- 
longs to  the  Natural  History  Museum 
(BMNH). 

Comments. — This  species  keys  out  to  M. 
inusitatum  (Snelling)  in  Michener  (1986), 
but  can  be  easily  distinguished  from  that 
species  by  the  malar  area  which  is  longer 
than  in  all  other  species  of  Mydrosoma ,  but 
linear  (absent)  in  M.  inusitatum.  It  might 
seem  to  belong  to  Friese's  Bicornelia  (sunk 
within  Mydrosoma  by  Michener  1986;  see 
also  Snelling  1980),  although  that  species 
group  is  defined  based  upon  secondary 
sexual  characteristics  of  males,  which  are 
unknown  for  M.  micheneri.  It  does  not  key 
out  to  either  species  of  "Bicornelia"  using 
Michener  (1986). 

The  locality  where  the  species  was 
collected  is  now  largely  agricultural,  al- 
though gallery  forest  remains  along  water- 
courses. There  is  a  substantial  area  of  forest 
to  the  west  of  the  type  locality  however,  the 


Parque  Nacional  do  Xingu.  It  is  possible 
that  this  species  may  persist  in  this  region. 

ACKNOWLEDGEMENTS 

I  am  grateful  to  George  Else  (BMNH)  for  the 
opportunity  to  borrow  the  specimen  described  herein. 
Jason  Gibbs  prepared  the  images  in  figures  1  and  2, 
for  which  I  am  grateful.  Funding  for  my  research  is 
provided  by  the  Natural  Science  and  Engineering 
Research  Council  of  Canada. 

LITERATURE  CITED 

Graf,  V.  and  D.  Urban.  2001.  Mydrosomella  cleia,  uma 
especie  nova  do  sul  do  Brasil  (Hymenoptera, 
Colletidae).  Acta  Biologica  Paranaense  30:  1715-179. 

Michener,  C.  D.  1986.  A  review  of  the  tribes 
Diphaglossini  and  Dissoglottini  (Hymenoptera: 
Colletidae).  University  of  Kansas  Science  Bulletin  53: 
183-214. 

— .  2007.  The  bees  of  the  world  [2"d  edition]. 
Johns  Hopkins  University  Press,  Baltimore,  Mary- 
land. 

Snelling,  R.  R.  1980.  The  genus  Bicornelia  (Hymenop- 
tera: Colletidae).  Contributions  in  Science,  Natural 
History  Museum  of  Los  Angeles  County  327:  1-6. 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  281-292 

Phenology  and  Social  Organization  of  Halictus  (Seladonia)  tripartitus 

(Hymenoptera:  Halictidae) 

Laurence  Packer,  Anne-Isabelle  D.  Gravel  and  Gretchen  Lebuhn 

(LP,  A-IDG)  Department  of  Biology,  York  University,  4700  Keele  St.,  Toronto, 

Ontario  M3J  1P3,  CANADA 
(GL)  Department  of  Biology,  San  Francisco  State  University,  San  Francisco,  CA,  94132,  USA 


Abstract. — We  present  data  on  the  social  biology  of  Halictus  (Seladonia)  tripartitus  Cockerell  based 
upon  samples  from  California.  This  bee  is  at  least  partially  eusocial  and  overwintered  gynes  and 
first  brood  workers  differed  in  size  by  5.14%.  35.5%  of  the  first  brood  workers  had  developed 
ovaries,  11.2%  had  ovaries  with  the  equivalent  of  at  least  one  fully  developed  oocyte  but  less  than 
14%  were  mated.  In  July  and  August,  female  reproductive  options  seemed  highly  variable: 
approximately  55%  were  sterile  workers,  2.6%  had  better  developed  ovaries  than  spring 
foundresses  and  over  20%  had  at  least  the  equivalent  of  one  fully  developed  oocyte  whereas  an 
additional  20-30%  may  have  been  capable  of  overwintering  as  gynes.  Ovarially  developed  workers 
were  larger  than  those  that  remained  sterile.  Sixteen  to  28%  of  the  late  summer  workers  were 
mated,  but  mated  individuals  were  not  more  likely  to  have  developed  ovaries  or  to  be  larger  than 
unmated  bees.  We  discuss  these  findings  in  the  light  of  the  climate  of  the  study  area  and  compare 
this  species  with  other  members  of  its  subgenus. 


Detailed  assessment  of  the  social  organi- 
sation of  halictine  bees  requires  painstak- 
ing field  observations  and  nest  excavations 
over  the  course  of  several  flight  seasons. 
Nonetheless,  useful  information  can  be 
obtained  from  less  detailed  analyses.  Be- 
cause of  the  evolutionary  lability  of  social- 
ity in  these  bees,  even  comparatively 
superficial  data  can  be  of  utility  in  terms 
of  plotting  presence  or  absence  of  sociality 
upon  a  phylogeny  or  in  obtaining  estimates 
of  sociobiologically  important  variables 
such  as  levels  of  morphological  and  phys- 
iological caste  differentiation  (e.g.  Dunn  et 
al.  1998). 

The  subgenus  Seladonia  (of  the  genus 
Halictus)  contains  species  that  are  primarily 
solitary,  such  as  H.  (S.)  virgatellus  Cockerell 
(Eickwort  et  al.  1996),  some  with  weak 
eusociality  (Michener's  1974  terminology 
for  social  categories  is  used  herein)  as  in  H. 
confnsus  Smith  (Dolphin  1971)  to  others 
with  the  largest  colony  sizes  ever  re- 
corded for  a  halictine  -  H.   (S.)  lutescens 


Friese  (Sakagami  and  Okazawa  1985)  or 
the  largest  morphological  caste  differenti- 
ation -  H.  (S.)  land  Moure  (Janjic  and 
Packer  2001).  However,  it  should  be  cau- 
tioned that  data  for  these,  and  most  other 
species  in  the  subgenus,  are  mostly  frag- 
mentary. 

Halictus  (Seladonia)  tripartitus  Cockerell  is 
a  halictine  for  which  the  only  sociobiolog- 
ical  data  available  are  i)  the  surprising 
observation  that,  at  a  nest  aggregation, 
smoke  blown  down  one  nest  entrance 
came  up  out  of  the  other  entrances, 
demonstrating  that  the  nests  were  con- 
nected underground  (Eickwort  personal 
communication  1988)  and  ii)  a  statement 
that  the  species  is  solitary  (Amdam  et  al. 
2006).  It  is  a  somewhat  divergent  member 
of  the  subgenus,  forming  the  sister  group 
to  the  remainder  (Danforth  et  al.  1999), 
probably  along  with  two  rare  species,  H. 
harmonius  Sandhouse  and  H.  pinguismentus 
Janjic  and  Packer  (Janjic  and  Packer  2001, 
Janjic  and  Packer  unpublished  data). 


282  Journal  of  Hymenoptera  Research 

For    a    halictine    population    with    an  species  of  bee  (Leong  and  Thorp  1999),  the 

archetypal  eusocial  phenology  (and  indeed  bee  fauna  was  surveyed  using  a  recently 

other  annual  eusocial  insects  in  temperate  developed   standard   protocol   which   uti- 

climates),  only  mated  and  comparatively  lizes   a    mix   of   sweep-netting   and   pan 

unworn  females  are  expected  to  survive  trapping    (protocols    can    be    found    at 

the  winter.  In  spring  they  establish  nests,  <http://online.sfsu.edu/~beeplot.>).  Pan 

their   wings   and   mandibles   increase   in  traps  were  set  out  between  09:00  and  15:00 

wear  and  their  ovaries  become  well  de-  and  sweep-netting  was  done  for  one  hour 

veloped  as   they  produce  a  brood  com-  in    the    morning   and    one   hour    in    the 

posed  primarily  of  worker  females.  Indi-  afternoon  per  sampling  day. 

viduals  in  this  brood  are  usually  smaller  Some  samples  of  females  from  2003  were 

than  their  mother  and  have  reduced  levels  preserved  in  70%  alcohol  and  this  permit- 

of  ovarian  development  and   a   low  fre-  ted  dissection  for  sociobiological  data  (see 

quency  of  mating  (partly  dependent  upon  below).  All  samples  collected  in  2002  were 

the  availability  of  males  produced  in  the  pinned  so  only  phenological  data  could  be 

"worker"   brood).    Depending   upon   the  obtained  from  them. 

length   of   the   flight   season,   additional  Sociobiological  Data. — The  preserved  bees 

broods,    primarily   of   workers,   may   be  were   observed    under   a    microscope    to 

produced.  Towards  the  end  of  the  colony  evaluate  alar  and  mandibular  wear  (based 

cycle,  a  brood  composed  of  males  and  next  upon  the  right  hand  side),  measure  head 

year's   colony   foundresses   is   produced,  width,  and  dissect  females  for  insemina- 

Much  of  the  variation  in  these  parameters  tion  and  ovarian  development  status  fol- 

can,  to  some  extent,  be  estimated  from  field  lowing   slight   modifications   of  standard 

collected  samples  in  the  absence  of  nest  protocols    (Ordway    1965,    Abrams    and 

excavations   (Dunn   et   al.    1998).    In   this  Eickwort    1980).    Mandibular   wear   was 

paper  we  present  some  phenological  and  scored   on  a   scale   from   0  -  completely 

social  organisation  data  for  H.  tripartitus  unworn,  apex  sharp;  to  6  -  mandible  worn 

based  upon  field  samples,  in  order  to  add  away  to  the  base  of  the  subapical  tooth, 

to  the  intensity  of  taxonomic  sampling  for  Wing  wear  was  assessed  by  counting  the 

such  data  in  these  behaviourally  diverse  number   of   nicks    in   the   wing   margin, 

bees.  completely  abraded  wings  were  scored  as 

having  15  nicks  -  the  number  that  seems  to 

MATERIALS  AND  METHODS  result  in  complete  abrasion  of  the  margin. 

Sampling. — Halictus    tripartitus    adults  Total  wear  was  calculated  by  adding  the 

were  collected  on  the  Kunde  Wine  Estate,  scores  for  wing  and  mandibular  wear.  It  is 

2.3  km  southeast  of  Kenwood  in  Sonoma  useful  to  exclude  freshly  emerged  adults 

County,    California,     USA     (38°24'15N,  from  some  analyses,  such  as  of  ovarian 

122  31'43W).  This  site  is  a  low-elevation,  development   and    mating,   because   they 

gently  sloping  oak  woodland  dominated  may  not  have  had  enough  time  to  mate 

by  blue  oak  (Quercus  douglasii),  adjacent  to  or  develop  their  ovaries.  Consequently,  in 

a  large  vineyard.  The  oak  woodland  con-  many  analyses  only  bees  with  a  total  wear 

sists  of  4-10  m  tall  trees  that  are  generally  score  greater  than  one  were  considered, 

widely   spaced   with   few    shrubs    inter-  Comparisons    of    ovarian    development 

spersed  among  them.  At  the  site,  a  100  m2  among  samples  were  performed  with  bees 

sampling  plot  was  established.  parasitized  by  conopids  removed  from  the 

Bees  were  sampled  in  2002  and  2003  on  sample.  In  later  samples,  when  a  mixture 
clear,  low  wind  days.  As  preliminary  data  of  ageing  workers  and  newly  emerged 
and  previous  monitoring  studies  show  that  gynes  might  be  expected,  putative  exam- 
different  colored  pan  traps  attract  different  pies  of  the  latter  were  assumed  to  have 


Volume  16,  Number  2,  2007 


283 


a  total  wear  of  at  most  one  and  to  have 
entirely  undeveloped  ovaries.  Individuals 
with  a  total  wear  of  three  or  more  and  with 
completely  undeveloped  ovaries  were  as- 
sumed to  be  permanently  sterile  workers 
in  these  samples.  This  protocol  seemed 
appropriate  in  this  study  as  these  bees 
appear  to  become  worn  rather  slowly,  their 
wings  seem  to  become  "nicked"  particu- 
larly slowly. 

The  spermatheca  of  each  bee  was  ob- 
served for  sperm,  which  make  the  other- 
wise glassy-transparent  spermatheca  opa- 
que. On  the  few  occasions  when  there  was 
uncertainty  as  to  whether  a  bee  had  mated 
or  not,  the  spermatheca  was  gradually 
squashed  between  a  cover  slip  and  a  mi- 
croscope slide  under  high  magnification  to 
look  for  spermatozoa.  Ovarian  develop- 
ment was  evaluated  by  estimating  the 
proportion  of  a  fully  developed  oocyte 
present  in  the  bees  and  summing  them 
across  the  6  ovarioles.  Bees  with  developed 
ovaries  (defined  as  with  at  least  one 
ovariole  with  at  least  one  quarter  of  a  fully 
developed  oocyte)  are  sometimes  referred 
to  as  OD+  in  the  account  that  follows,  bees 
without  ovarian  development  are  referred 
to  as  OD  — .  Caste  size  dimorphism  is 
estimated  as  (q-w)/q  where  q  is  mean 
putative  queen  head  width  and  w  is  mean 
putative  worker  head  width. 

Climatic  data. — Because  weather  varia- 
tion, both  within  and  among  years,  influ- 
ences sociobiological  parameters  for  sweat 
bees  (Richards  and  Packer  1995),  we  report 
temperature  and  rainfall  data  for  the  study 
area  both  in  terms  of  data  from  the  period 
when  bees  were  sampled  and  for  longer 
term  average  conditions.  Data  were  taken 
from  the  Santa  Rosa  Airport  Weather 
Station  as  indicated  on  the  following  web- 
pages:  http://www.pressdemocrat.com/ 
nbwx/srweather/index.cfm  and  http:// 
www.wunderground.com/history/airport/ 
KSTS/1993/3/l/MonthlyHistory.html# 
calendar.  Data  presented  here  are  for  mean 
daily  temperature  and  rainfall  for  the 
months  of  March  through  August  2002  and 


300  -i 

r  18 

co250" 
®  200  - 

w 

\  \ 
\  \ 

-  16 

-  14 

-  12 

CO 

E   150  - 

/ 
/ 

t               1 

\   \ 
\    \ 

-  10 

-  8 

£  ioo- 

^ 

-  6 

-  4 

50  - 
0  - 

• 

-  2 

1 

1                          V 

May 

Jun  Jul 

Aug  Sep  Oct 

(f) 

CO 


* 


Fig.  1.  Phenology  of  H.  tripartitus.  The  left-hand  axis 
indicates  the  number  of  females  (•)  while  the  right  axis 
shows  the  number  of  males  (A),  all  collected  in  2002. 

2003,  along  with  averages  and  maxima  and 
minima  for  the  same  months  over  the 
10  year  period  1992-2001. 

RESULTS 

Phenological  Patterns. — During  the  May- 
September  period,  2002,  a  total  of  765  H. 
tripartitus  were  collected  at  the  Kunde 
Wine  Estate.  Of  these,  726  were  females 
and  39  were  males.  Numbers  of  H.  triparti- 
tus females  increased  gradually,  nearly  at 
a  constant  rate,  from  May  to  mid-July 
(Fig.  1)  peaking  at  the  beginning  of  August 
after  a  temporary  decrease  in  sample  size, 
and  then  decreased  rapidly  until  the  end  of 
the  season.  Males  show  a  similar  pattern, 
but  peak  in  the  sampling  period  after  the 
females.  The  discrepancy  between  the 
number  of  males  and  number  of  females 
in  the  samples  is  surprising;  even  at  the 
peak  of  male  abundance,  there  are  approx- 
imately 8  times  as  many  females  as  males. 
Based  upon  observations  of  museum  hold- 
ings, this  ratio  would  seem  not  to  be  an 
artefact  as  males  of  H.  tripartitus  are  rare  in 
collections  (Packer  unpublished  observa- 
tions). 

The  alcohol  preserved  material  demon- 
strates that  the  species  is  active  as  early  as 
March  (see  below),  and  inspection  of 
museum  holdings  shows  that  a  few  fe- 
males can  be  found  as  late  as  October 
(Packer  unpublished  observations). 

The  March  28,h  sample  of  dissected  bees 
suggest  that  nest  initiation  had  only  just 
begun  at  this  time:  only  39%  of  the  bees 


284 


Journal  of  Hymenoptera  Research 


3.5 

3 

A 

A 

c    2.5 

0) 

E 

Q. 

o       2 

0) 

> 

?    15- 

A 

A                             A 

A                           A 

A        A 

A                   A 

A 

*                 k                 A 

A      A 

(0 

1  1 

*        A 

A 

^        A                  ' 

A 

A 

0.5 

A 

U  1 — 
1.8 

i 

1.9 

1                                                         * 

2                    2.1 

2.2 

2.3 

Head  width  (mm) 

Fig.  2.     Ovarian  development  and  size  variation  in  females  of  the  May  sample. 


had  mandibular  wear,  16%  had  wing  wear 
and  34%  of  them  had  some  ovarian  de- 
velopment with  mean  summed  oocyte 
fractions  of  0.16  per  individual  (Table  1). 
There  was  no  size  difference  between 
ovarially  developed  and  undeveloped  fe- 
males (mean  head  width  OD+  females  = 
2.04  mm,  SD  =  0.11,  n  =  54;  OD-  females 
head  width  =  2.06  mm,  SD  =  .093,  n=  28;  t 
=  0.93,  ns).  Almost  all  females  mated 
(97.2%),  both  unmated  females  were  above 
average  size  for  the  sample. 


In  contrast,  the  May  16th  sample  was 
entirely  of  mated  bees  with  worn  mand- 
ibles, 94%  had  worn  wings  and  all  except 
a  parasitized  individual  had  developed 
ovaries  with  the  summed  fractions  of 
developing  oocytes  averaging  over  1.6  per 
bee.  Ovarian  development  was  indepen- 
dent of  bee  size  (Fig.  2).  The  mean  size  of 
March  and  May  sample  bees  was  2.04  (SD 
=  0.10)  and  2.05  mm  (SD  =  0.09)  respec- 
tively, not  significantly  different  (t  =  0.35, 
p  >  0.5). 


Table  1.     Summary  sociobiological  data  by  sample  date. 


%  of  9  with 

%  of  9  with 

".i  of  9  with* 

Mean  ovarian 

Mean  9  Head 

Sample  date 

N 

mandible  wear 

wing  wear 

%  of  9  mated 

developed  ovaries 

score* 

width 

March  28 

82 

39 

16 

97.2 

34 

0.16 

2.04 

May  16 

31 

100 

94 

100 

100 

1.6 

2.05 

June  20 

67 

81.5 

26 

13.8 

43 

0.4 

1.93 

June  30 

50 

96 

62 

6.5 

26 

0.14 

1.94 

July  18 

166 

68.7 

70.5 

13.7 

55 

0.44 

2.01 

August  20 

118 

56 

36.2 

34.2 

34.5 

0.28 

2.01 

I  fema  riitted  from  ovarian  development  data. 


Volume  16,  Number  2,  2007 


285 


The  two  samples  from  June  show  an 
increased  variance  in  wear  but  decreased 
ovarian  development  in  comparison  to  the 
May  sample.  On  June  20*  81.5%  of  the 
bees  had  worn  mandibles,  26%  had  worn 
wings,  43%  had  developed  ovaries  and 
13.8%  were  mated.  This  was  the  only 
sample  in  which  unworn  bees  were  signif- 
icantly less  likely  to  have  developed 
ovaries  (x2  =  7.18,  P  <  0.01;  all  other 
samples  p  >  0.16),  suggesting  that  many  of 
these  individuals  were  young  and  had  not 
had  time  to  develop  their  ovaries.  None- 
theless, worn  bees  with  undeveloped  ova- 
ries made  up  46.5%  of  this  sample.  On  June 
30th  comparable  data  are:  96%  with  worn 
mandibles,  62%  with  worn  wings,  26% 
with  developed  ovaries  (31%  if  only  worn 
bees  are  included)  and  6.5%  were  mated. 
These  data  suggest  increased  average  age/ 
activity  levels  of  the  bees  over  the  in- 
tervening ten  days  but  a  decrease  in 
ovarian  development.  The  average  sum  of 
fractions  of  oocytes  shows  the  same  de- 
crease over  time,  the  values  were  0.40  for 
[une  20th  and  0.14  for  June  30th.  Of  all  June 
bees  combined,  only  one  individual  (ie  < 
1%)  had  more  enlarged  ovaries  than  the 
average  individual  in  the  May  sample. 

Ovarially  developed  worn  bees  were 
significantly  larger  than  worn  OD-  bees 
in  the  June  30th  sample  (mean  head  width 
OD+  bees  =  1.98  mm,  SD  =  0.076,  n  -  11; 
OD-  =  1.92  mm,  SD  =  .089,  n  =  28;  t  = 
2.1,  p<  0.05),  but  not  in  the  June  20th  one 
(OD+  =  1.95  mm,  SD  =  0.94,  n  =  23,  OD- 
=  1.93  mm,  SD  =  .096,  n  =  19;  t  =  0.91,  ns). 
The  proportion  of  bees  that  were  mated 
did  not  differ  between  the  two  June 
samples  (Fisher's  exact  test,  p  =  0.35). 

The  data  suggest  that  these  early  sum- 
mer bees  are  offspring  of  the  generation 
sampled  in  March  and  May  and  that  they 
represent  a  worker  brood.  The  pattern  of 
ovarian  development  and  age  in  these  bees 
suggests  that  more  workers  initiate  ovarian 
development  soon  after  eclosion  than 
manage  to  maintain  developed  ovaries 
a  short  while  later.  Furthermore,  the  data 


indicate  that  individuals  that  do  maintain 
developed  ovaries,  are  disproportionately 
larger  than  those  that  do  not. 

Bees  in  the  two  June  samples  did  not 
differ  in  size  (1.93  mm  and  1.94  mm  for 
June  20th  and  June  30th  respectively).  Three 
individuals  collected  in  June  were  extreme- 
ly worn,  (wing  margin  completely  abrad- 
ed), suggesting  that  they  may  be  ageing 
overwintered  females,  perhaps  from  mul- 
tiple foundress  associations.  All  three  were 
mated,  one  was  parasitized  with  a  conopid 
larva  (Diptera,  Conopidae),  one  had  no 
ovarian  development,  the  remaining  in- 
dividual had  Va  of  a  developed  oocyte. 
Inclusion  of  these  individuals  in  the  com- 
parison of  ovarian  development  data  be- 
tween worn  and  unworn  bees  did  not  alter 
the  statistical  patterns  noted  above. 

The  July  18th  sample  had  a  lower  pro- 
portion of  bees  with  mandibular  wear  than 
in  June  (68.7%),  a  higher  proportion  with 
worn  wings  (70.5%)  and  a  higher  pro- 
portion with  developed  ovaries  (55%, 
54.4%  when  only  worn  bees  are  included). 
Similar  to  the  June  20th  sample,  13.7%  of 
females  were  mated.  The  wear  differential 
between  mandibles  and  wings  suggests 
that  bees  at  this  time  are  performing 
relatively  more  foraging  or  less  digging 
than  in  the  earlier  samples.  The  average 
sum  of  fractions  of  oocytes  was  0.44  and  6 
individuals  (3.8%)  had  more  ovarian  de- 
velopment than  the  average  overwintered 
female  in  May.  Worn  OD+  bees  were 
significantly  larger  than  worn  OD-  bees 
(head  width  OD+  =  2.02  mm,  SD  =  0.07,  n 
=  82;  head  width  OD-  -  1.98  mm,  SD  = 
0.09,  n  =  44  respectively;  t  =  2.56,  p  < 
0.05).  Overall,  July  bees  were  intermediate 
in  size  between  the  smaller  June  and  larger 
earlier  samples,  averaging  2.01  mm  in 
head  width;  the  sample  of  six  individuals 
with  very  well  developed  ovaries  had 
exactly  the  same  average  head  width  as 
the  rest  of  the  sample. 

On  August  20th  the  proportion  of  bees 
with  worn  mandibles  decreased  to  56% 
while  36.2%  had  worn  wings.  The  pro- 


286 


Journal  of  Hymenoptera  Research 


Wear  and  Reproductive  Data  for  Different  Sample  Dates 

q         <y 

worn  mandibles  -  -n-  -  %  worn  wings  -  -a-  -  %  ovarially  developed      -♦-      %  mated 

120  - 
mn 

■~~  < 

80 

X/       \\                \ 

ntage 

o 

/y  /               *\       P~       ^^^, 

CD 

u 

//         '                                                \      \                  '                    A^             ^            ^^3 

Q. 

40  < 

4 

20  - 

/ 

[ 

J 

u 

0 

20                 40                 60                 80                100               120               140               160 

Day:  March  28th  =  day  1 

Fig.  3.     Wear  and  reproductive  variables  for  each  of  the  samples  of  bees  dissected. 


portion  with  developed  ovaries  decreased 
to  34.5%  (38.5%  when  only  worn  bees  are 
included)  and  the  percentage  of  females 
that  had  mated  increased  to  34.2%.  The 
summed  fraction  of  oocytes  declined  to  an 
average  of  0.28  and  only  one  individual 
(<1%  of  the  total)  had  ovaries  that  ex- 
ceeded the  average  ovarian  development 
of  the  May  sample.  As  with  the  July 
sample,  worn  OD+  bees  in  August  were 
significantly  larger,  on  average,  than  OD— 
individuals  (head  width  OD+  =  2.04  mm, 
SD  =  =  0.10,  n  ^  38;  head  width  OD- 
1.99  mm,  SD  =  0.12,  n  =  28  respectively;  t 
2.73,  p  <  0.01).  Overall,  the  bees 
collected  in  August  had  an  average  head 
width  of  2.01  mm,  the  individual  with  very 
well  developed  ovaries  had  a  head  width 
of  2.07  mm  (not  significantly  larger  than 
the  sample  as  a  whole,  Mann  Whitney  U 
test,  p  =  0.65). 

The  wear   and   reproductive   data   dis- 
cussed above  are  shown  visually  in  Fig.  3. 


Size  variation  among  the  four  samples 
(March  and  May  samples  combined,  two 
June  samples  combined)  is  significant 
(ANOVA,  F  =  6.0,  p  =  0.0005).  Tukey's 
HSD  test  reveals  that  this  result  is  due  to 
the  June  bees  being  significantly  smaller 
than  the  others  (p<0.01  for  the  overwin- 
tered female  comparison  and  p<0.01  and 
p<  0.05  for  the  comparisons  with  July  and 
August  samples  respectively). 

Caste. — The  data  above  are  consistent 
with  the  March  sample  consisting  of  over- 
wintered females  at  the  nest  initiation 
phase,  the  May  sample  comprising  the 
same  generation  individuals  actively  pro- 
visioning nests  and  with  the  June  sample 
being  first  brood  workers.  The  three 
heavily  worn  June  bees  perhaps  represent 
subordinates  in  multiple  foundress  asso- 
ciations. Although  this  would  require  nest 
excavation  data  for  confirmation,  it  is 
uncommon  for  solitary  foundresses  to 
forage  for  such  an  extensive  period  of  time 


Volume  16,  Number  2,  2007 


287 


Overwintered  females  (March  +  May) 


30 
20 

10 
0 


40 
30 
20 
10 
0 


20 
15 

10  -I 
5 
0 


1st  workers  (June) 


...lllllIlM 


July 


^Jl 


August 


■    ■    l.lllllll- 


1.6     1.67    1.73     1.8     1.87    1.93      2      2.07    2.13     2.2     2.27 
Head  width  (mm) 


"ig.  4.  Size  variation  (head  width)  among  females 
rom  each  of  the  months  sampled. 

3-r  to  become  so  heavily  worn.  If  this  is  the 
:ase,  then  the  putative  gynes  average 
2.04  mm  in  head  width  (SD  =  0.10)  and 
:he  first  worker  brood  1.94  mm  (SD  = 
3.09),  the  castes  differ  significantly  in  size  (t 
=  11.22,  p  <<  0.001)  and  the  morpholog- 
cal  size  difference  between  gynes  and  the 
:irst  worker  brood  is  5.14%. 

Size  variation  data  for  the  overwintered 
:emales,  first  brood  workers  and  July  and 
August  females  are  shown  in  Fig.  4. 


July  and  August  females  are  not  so  easily 
categorised  as  to  caste  as  they  are  expected 
to  include  some  newly  emerged  gynes  as 
well  as  ageing  workers  and  perhaps  some 
young  workers  also.  We  take  ageing  bees 
with  undeveloped  ovaries  to  be  sterile 
workers,  ageing  bees  with  well  developed 
ovaries  (with  total  ovarian  development  at 
least  equivalent  to  one  fully  developed 
oocyte)  to  be  potentially  reproductive 
workers  and  unworn  bees  with  completely 
undeveloped  oocytes  to  potentially  be 
gynes. 

Data  for  these  three  groups  are  shown  in 
Table  2  separately  for  July  and  August. 
Interestingly,  the  sizes  of  the  potential 
gynes  in  July  are  very  similar  to  those  for 
the  overwintered  females  the  previous 
March  and  May.  The  reproductive  workers 
are  also  somewhat  large,  but  their  wear 
indices  suggest  that  they  do  not  represent 
the  same  overwintered  females  as  were 
sampled  in  March  and  May  as  their  index 
of  wear  is  too  low  for  them  to  have  been 
active  for  the  entire  intervening  period 
(only  three  of  40  have  a  higher  index  of 
wear  than  the  average  female  in  the  May 
sample)  and  none  of  them  are  mated.  The 
sterile  workers  are  the  smallest  individuals 
in  the  sample. 

The  pattern  for  the  same  three  classes  of 
bee  for  the  August  sample  is  quite  differ- 
ent. In  August  the  sterile  worker  and 
potential  gyne  samples  are  identical  in  size 
but  the  reproductive  workers  are  larger. 
Indeed,  the  mean  size  of  the  reproductive 
workers  in  August  is  the  largest  of  any  of 
the  categories  analysed  in  any  time  period. 
Even  these,  however,  would  not  seem  to  be 
remaining  overwintered  females  as  only  4 


Table  2.     Sociobiological  data  for  different  "castes"  of  summer  female.  For  explanation  of  caste  designation 
;ee  text. 


July 

August 

Putative  Caste 

Head  width  (SD)  N 

%  mated 

Head  width  (SD)  N 

"..  mated 

sterile  workers 
Reproducing  workers 
3ynes 

1.99  (0.07)  22 
2.02  (0.09)  5 
1.99  (0.09)  30 

13.6 

0.0 

28.9 

1.99  (0.09)   10 

2.10  (0.14)  5 
2.03  (0.08)  45 

60.0 

0.0 
17.4 

288 


Journal  of  Hymenoptera  Research 


of  the  15  have  a  total  wear  greater  than  the 
May  sample  and  none  of  them  are  mated. 

It  would  seem  that  the  fates  of  bees 
eclosing  in  summer  are  highly  variable. 
Some  seem  to  remain  strictly  as  workers, 
attaining  high  wear  indices  without  de- 
veloping their  ovaries.  A  total  of  55%  of  the 
July  and  August  bees  had  a  total  wear 
index  of  3  or  more  but  had  completely 
undeveloped  ovaries.  Such  bees  were 
smaller  than  the  worn  bees  that  had 
developed  ovaries  and  the  size  difference 
between  these  apparent  permanent  work- 
ers and  their  contemporaries  that  have 
highly  developed  ovaries,  with  ovarian 
development  greater  than  or  equal  to  one 
is  even  larger:  3.8%  (t  =  5.24,  p  <<  0.001) 
and  4.8%  (t  =  4.54,  p  <<  0.001)  for  July 
and  August  samples  respectively. 

For  none  of  the  samples  of  worn  bees 
was  there  a  significant  association  between 
ovarian  development  and  mating  (p  >  0.5 
in  all  comparisons).  Similarly,  the  mated 
bees  were  not  larger  than  unmated  ones 
(worn  bees  only)  in  any  of  the  analyses  (p 
>  0.5  in  each  case)  except  for  the  July 
sample  in  which  the  mated  bees  were 
significantly  larger  than  the  unmated  ones 
(head  width  mated  =  2.05  mm,  SD  =  0.08, 
n  =  14,  head  width  unmated  =  1.99  mm, 
SD  =  0.06,  n  -  75;  t  =  7.14,  p  <<  0.001). 

Parasitism. — One  female  was  found  with 
a  large  nematode  in  the  metasoma,  she  was 
from  the  March  sample  of  overwintered 
females.  Twenty  females  were  parasitized 
with  one  conopid  larva  in  each  metasoma 
and  three  contained  two  parasite  larvae.  It 
is  likely  that  these  parasitism  rates  are 
underestimates  as  small  conopid  larvae 
and  nematodes  would  likely  have  been 
missed  in  the  dissections,  especially  if  they 
were  primarily  underneath  the  first  meta- 
somal  tergum,  which  was  not  removed. 

Of  the  23  females  with  conopids,  one 
was  found  in  the  May  sample,  four  each  in 
the  two  June  samples,  eight  in  July  and  six 
in  August.  The  overwintered  females  were 
significantly  less  affected  by  conopids  than 
were  later  females  (Fisher's  exact  test  p  = 


0.0377).  The  long  period  during  which 
conopid  larvae  were  found  suggests  either 
that  the  parasite  has  more  than  one 
generation  per  year,  or  that  multiple 
species  of  conopid  are  involved. 

Bees  with  conopid  larvae  did  not  differ 
in  size  from  the  other  individuals  in  their 
samples  (p  >0.2  in  all  cases),  but  they  did 
have  more  wear  than  their  contemporaries 
in  July  and  August  (Mann  Whitney  U  test, 
U  =  991,  p  =  0.007  and  U  =  536,  p  =  0.014 
respectively)  but  not  in  either  of  the  June 
samples  (U  =  191,  p  =  0.059  and  U  =  130, 
p  =  0.187). 

Climate  and  Weather. — The  collection  pe- 
riod for  H.  tripartitus  was  from  May  to 
August  2002  for  the  pinned  specimens  and 
from  March  to  August  2003  for  the  pickled 
ones.  The  weather  data  for  the  months 
March  to  August  are  shown  in  Fig.  5  along 
with  averages  for  the  previous  ten  year 
period.  2002  was  drier,  but  2003  was  wetter 
than  the  average  for  the  previous  ten  year 
period.  In  both  years  April  was  an  extreme 
month,  with  more  than  double  the  average 
rainfall  in  2003  and  less  than  a  quarter  the 
normal  amount  in  2002.  The  temperature 
data  show  that  2002  varied  little  from  the 
average  conditions  whereas  in  2003  April 
was  cooler  than  any  year  between  1992  and 
2002  and  July  was  warmer  than  in  any  of 
these  other  years. 

DISCUSSION 

Halictine  bees  are  well  known  for  their 
variable  social  behaviour,  not  only  within 
and  between  populations  and  species  but 
also  among  individuals  within  a  colony. 
Taken  in  their  entirety,  our  data  strongly 
support  the  view  that  H.  tripartitus  is 
primarily  eusocial  at  our  study  site.  Per- 
haps the  strongest  evidence  for  this  comes 
from  all  overwintered  females  being  mated 
whereas  the  first  brood  offspring  were 
almost  entirely  unmated.  Indeed,  as 
a  whole,  the  data  are  mostly  consistent 
with  the  view  that  this  species  is  a  typical 
annual  eusocial  species  with  overwintered 
females    initiating   nests    in    March    and 


Volume  16,  Number  2,  2007 


289 


35 


30 


25 


20 


O 

(0 

2    15 

£ 
fi    10 


0 


march 


april 


may 


june 


July 


august 


-  -  •-  -  average 


lowest 


highest  -  -a-  -  2002       a      2003 


5  - 


1    4 


JS    3 


a  D 


_i 


i 


march 


april 


may 


lune 


July 


august 


2003  □  average  0  2002 


Fig.  5.     Weather  data  for  2002,  2003  and  summaries  for  the  ten  previous  years  1992-2001. 


foraging  until  May,  workers  emerging  in 
June  and  overwintering  gynes  commenc- 
ing emergence  in  July.  Less  usual  features 
are  the  apparent  multiple  nest  entrances, 
low  frequency  of  males  and  the  nature  of 
the  sample  in  August.  We  return  to  these 
three  aspects  towards  the  end  of  the 
discussion  after  considering  worker  fitness 
options  and  making  comparisons  between 


H.    tripartitus   and    other   species   of   the 
subgenus  Seladonia. 

Fitness  options  for  bees  eclosing  in 
summer  within  a  eusocial  family  structure 
include  remaining  as  a  sterile  worker, 
attempting  to  produce  some  offspring  di- 
rectly or  initiating  a  nest  either  the  same 
year  as  they  eclose  (Richards  et  al.  2003)  or 
after  overwintering  (Yanega  1988).  Given 


290 


Journal  of  Hymenoptera  Research 


the  Mediterranean  climate  of  the  region 
and  the  dryness  and  hardness  of  the  soil  in 
summer,  initiating  a  nest  in  summer  is 
probably  not  a  very  profitable  option 
(McCorquodale  1989).  Furthermore,  as  the 
most  ovarially  developed  summer  bees 
were  unmated,  it  is  unlikely  that  they  are 
founding  new  nests  at  this  stage,  unlike 
some  summer  females  of  Halictus  (Halictus) 
sexcinctus  (Richards  et  al.  2003). 

Halictus  tripartitus  worker  brood  individ- 
uals seem  to  take  advantage  of  at  least 
three  of  these  options.  Of  the  July  and 
August  samples,  approximately  55%  seem 
to  be  sterile  workers  and  perhaps  20%  of 
them  seem  capable  of  reproducing.  Over 
2%  of  the  bees  collected  in  July  and  August 
had  ovaries  more  fully  developed  than  did 
the  foundresses  in  spring,  suggesting  that 
either  some  are  nesting  solitarily  or  they 
have  the  potential  to  reproduce  in  their 
natal  colony  (as  has  been  demonstrated 
using  genetic  markers  in  several  other 
species,  Packer  and  Owen  1994,  Richards 
et  al.  1995).  As  none  of  these  most  ovarially 
developed  later  workers  were  mated,  the 
offspring  they  produce  must  be  male. 
However,  few  males  are  ever  seen  in  this 
species.  This  would  be  in  agreement  with 
Packer  and  Owen  (1994)  where  genetic 
data  showed  that  considerable  levels  of 
ovarian  development  resulted  in  few  in- 
stances of  successful  oviposition  by  work- 
ers of  Lnsioglossum  laevissimum. 

The  fourth  option,  mated  worker  brood 
females  overwintering  and  initiating  a  nest 
the  following  spring,  cannot  be  confirmed 
or  refuted  with  the  data  at  hand.  It  remains 
possible  that  brood  divalency  may  occur  in 
H.  tripartitus.  The  easiest  way  to  document 
this  would  be  to  mark  large  numbers  of 
emerging  "worker"  brood  females  as  they 
leave  the  nest  and  search  for  them  when 
the  foundresses  begin  activity  the  follow- 
ing spring. 

Few  species  of  the  subgenus  Seladonia 
have  received  detailed  sociobiological  in- 
vestigation. The  best  studied  is  H.  (S.) 
hesperus  Smith,  which  has  large  colonies 


and  very  large  morphological  caste  differ- 
entiation with  concomitantly  low  worker 
ovarian  development  (<1%  of  workers 
seemingly  capable  of  reproduction,  Brooks 
and  Roubik  1983,  Packer  1985).  Halictus  (S.) 
lutescens  has  even  larger  colony  sizes  and 
in  the  one  nest  excavated  by  Sakagami  and 
Okazawa  (1985)  approximately  one  quarter 
of  the  workers  seemed  capable  of  repro- 
duction, though  it  remains  likely  that  this 
colony  was  studied  after  the  death  of  the 
queen.  A  third  species,  H.  (S.)  lanei,  has  the 
largest  caste  size  dimorphism  of  any 
halictine  known  (Janjic  and  Packer  2001), 
although  its  social  organisation  may  not  fit 
the  standard  eusocial  model  (Gravel  et  al. 
in  preparation).  All  three  of  these  species 
are  tropical  and  their  workers  seem  not  to 
have  the  range  of  options  that  may  be 
available  to  H.  tripartitus.  In  contrast,  the 
alpine  H.  (S.)  virgatellus  is  predominantly 
solitary  with  a  few  nests  being  shared 
(Eickwort  et  al.  1996).  The  north  temperate 
H.  (S.)  confusus  and  its  sibling  species  H. 
(S.)  tumulorum  (L.)  are  weakly  eusocial, 
perhaps  reverting  to  solitary  behaviour  at 
the  northern  edge  of  their  ranges  (Dolphin 
1971,  Sakagami  and  Ebmer  1979).  Worker 
brood  individuals  in  these  species  likely 
have  a  similar  range  of  options  as  sug- 
gested above  for  H.  tripartitus,  albeit 
usually  within  a  shorter  summer  activity 
period,  but  appropriately  detailed  studies 
remain  to  be  published. 

Richards  and  Packer  (1995)  found  that 
variations  in  local  weather  patterns  affect- 
ed the  demography  and  behaviour  of  H. 
(H.)  ligatus  colonies:  warmer  and  drier 
weather  resulted  in  larger  broods  with 
better  survival  rates  and  lower  nest-failure 
and  more  reproduction  by  workers.  Thus, 
warm,  dry  weather  resulted  in  weaker 
eusociality.  Conversely,  colder,  wetter  con- 
ditions gave  rise  to  smaller  workers  with 
greater  reproductive  differentiation  be- 
tween the  castes  but  also  lower  brood- 
survival  rates  and  more  nest-failure. 
Colder  and  wetter  weather  resulted  in 
stronger  eusociality.   The  area   where  H. 


Volume  16,  Number  2,  2007 


291 


tripartitus  was  studied  was  unusually  cold 
and  wet  in  April  of  2003,  during  foundress 
provisioning.  If  the  weather  influences  this 
species  in  the  same  manner  as  it  does  with 
H.  ligatus,  it  is  likely  that  early  summer 
colonies  were  smaller,  with  stronger  phys- 
iological caste  differentiation  than  is  usual 
for  this  population.  The  poor  weather  in 
April  may  also  explain  the  apparently 
extended  period  of  activity  of  overwin- 
tered foundresses. 

There  are  no  detailed  studies  of  social 
sweat  bees  from  Mediterranean  climates  in 
North  America.  The  restriction  of  precipi- 
tation to  the  winter  and  early  spring  is 
a  feature  that  can  limit  the  duration  of 
colonies  of  social  bees  in  the  area  and  few 
individuals  of  H.  tripartitus  have  been 
collected  from  late  August  onwards:  even 
though  temperatures  are  adequate  for 
activity,  the  dry  conditions  severely  re- 
stricts the  availability  of  forage.  It  is  likely 
that  the  late  summer  activity  of  this 
species  is  a  comparatively  recent  phenom- 
enon as  at  this  time  of  year  they  rely  upon 
flowering  of  plants  that  are  maintained 
by  agricultural  irrigation.  This  could  ex- 
plain the  unusual  nature  of  the  August 
sample. 

Two  other  unusual  aspects  of  this  spe- 
cies' biology  are  worthy  of  comment,  and 
may  be  related.  First,  the  very  low  pro- 
portion of  males,  either  as  sampled  here  or 
in  museum  collections,  is  surprising.  To- 
wards the  end  of  the  colony  cycle  in  most 
eusocial  halictines,  males  are  approximate- 
ly as  common  as  females,  yet  in  H. 
tripartitus  they  are  almost  an  order  of 
magnitude  less  common.  Second,  the  ob- 
servation of  nests  apparently  being  con- 
nected underground  is  entirely  unique 
among  bees.  It  remains  possible  that  males 
primarily  search  for  mates  underground, 
thereby  uniting  these  two  unusual  phe- 
nomena. Certainly  this  species  is  worthy  of 
more  detailed  field  investigations,  some- 
thing we  hope  this  paper  will  stimulate. 

Lastly,  the  observation  that  worker  ovar- 
ian development  seemed  to  decline  over 


time  is  of  interest.  This  phenomenon  has 
been  observed  in  numerous  social  insects. 
There  are  two  main  reasons  this  might  be 
the  case.  First,  the  act  of  work  might  make 
it  less  likely  that  a  bee  has  the  energy 
resources  to  develop  oocytes  -  which  are 
remarkably  large  in  comparison  to  bee 
body  size  in  halictines.  Alternatively,  it 
could  be  that  workers  develop  their  ovaries 
primarily  to  coincide  with  male  production 
when  reproductive  broods  are  protan- 
drous.  The  fact  that  first  brood  workers 
are  active  at  a  time  of  year  when  few  males 
are  being  produced  in  H.  tripartitus,  argues 
against  the  second  explanation. 

AKNOWLEDGEMENTS 

The  senior  author's  research  is  funded  by  the 
Natural  Sciences  and  Engineering  Research  Council 
of  Canada.  We  are  grateful  for  the  comments  of 
Miriam  Richards  on  an  earlier  version  of  this 
manuscript  and  to  Luana  Sciullo  for  processing  the 
weather  data.  The  junior  author's  research  is  funded 
by  the  Integrated  Hardwoods  Range  Management 
Program.  We  also  thank  Cynthia  Fenter,  Erin  Rentz 
and  many  SFSU  students  for  their  work  collecting 
these  insects. 

LITERATURE  CITED 

Abrams,  J.  and  G.  C.  Eickwort.  1980.  Biology  of  the 
communal  sweat  bee,  Agapostemon  virescens  (Hy- 
menoptera:  Halictidae)  in  New  York  state.  Search 
(Cornell  University  Agriculture  Experiment  Station) 
1:  1-20. 

Amdam,  G.  V.,  A.  Csondes,  M.  K.  Fondrk,  and  R.  E. 
Page,  Jr.  2006.  Complex  social  behaviour  derived 
from  maternal  reproductive  traits.  Nature  439: 
76-78.  [Cover  photo  caption  associated  with 
article.] 

Brooks,  R.  W.  and  D.  W.  Roubik.  1983.  A  Halictine  bee 
with  distinct  castes:  Halictus  hesperus  (Hymenop- 
tera:  Halictidae)  and  its  bionomics  in  Central 
Panama.  Sociobiology  7:  263-282. 

Danforth,  B.  N.  2002.  Evolution  of  sociality  in 
a  primitively  eusocial  lineage  of  bees.  Proceedings 
of  the  Natural  Academy  of  Science  of  the  United  States 
of  America  99:  286-290. 

,  H.  Sauquet,  and  L.  Packer.  1999.  Phytogeny  of 

the  bee  genus  Halictus  (Hymenoptera:  Halictidae) 
based  on  parsimony  and  likelihood  analyses  of 
nuclear  EF-loc  sequence  data.  Molecular  Phyloge- 
netics  and  Evolution  13:  605-618. 

Dolphin,  R.  E.  1971.  Observations  of  Halictus  confusus 
Smith  (Hymenoptera:  Halictidae)  on  woodland 


292 


Journal  of  Hymenoptera  Research 


and  field  flowers.  Proceedings  of  the  Indiana 
Academy  of  Science  81:  182-185. 

Dunn,  M,  P.  L.  Mitchell,  and  L.  Packer.  1998. 
Phenology  and  social  biology  of  two  sibling 
species  of  Halictus  in  an  area  of  sympatry. 
Canadian  Journal  of  Zoology  76:  2207-2213. 

Eickwort,  G.  C,  J.  M.  Eickwort,  J.  Gordon,  and  M.  K. 
Eickwort.  1996.  Revision  to  solitary  behavior  from 
eusocial  ancestry  in  the  sweat  bee  Halictus  rubi- 
cundus  in  the  Rocky  Mountains,  and  its  implica- 
tions for  high-althtude  and  high-latitude  adapta- 
tions and  holarctic  distributions  of  social  insects. 
Behavioral  Ecology  and  Sociobiology  38:  227-233. 

Janjic,  J.  and  L.  Packer,  L.  2001.  New  descriptions  of 
Halictus  (Seladonia)  from  the  New  World  (Hyme- 
noptera: Halictidae).  journal  of  Hymenoptera  Re- 
search 10:  55-75. 

Leong,  J.  M.  and  R.  W.  Thorp.  1999.  Colour-coded 
sampling:  the  pan  trap  colour  preferences  of 
oligolectic  and  nonoligolectic  bees  associated 
with  a  vernal  pool  plant.  Ecological  Entomology 
24:  329-335. 

McCorquodale,  D.  B.  1998.  Soil  softness,  nest  initiation 
and  nest  sharing  in  the  wasps,  Cerceris  antipodes 
(Hymenoptera:  Sphecidae).  Ecological  Entomology 
14:  191-196. 

Michener,  C.  D.  1974.  The  Social  Behavior  of  Bees:  A 
Comparative  Study.  Harvard  University  Press, 
Cambridge,  MA. 

Ordway,  E.  1965.  Caste  differentiation  in  Augochlorella 
(Hymenoptera,  Halictidae).  Insectes  Sociaux  12: 
291-308. 


Packer,  L.  1985.  The  social  organisation  of  two 
halictine  bees  from  southern  Mexico  with  notes 
on  two  bee-hunting  philanthine  wasps.  Pan- 
Pacific  Entomologist  51:  291-298. 

and  R.  E.  Owen.  1994.  Relatedness  and  sex 

ratio   in   a   primitively   eusocial   halictine   bee. 
Behavioral  Ecology  and  Sociobiology  34:  1-10. 

Richards,  M.  H.  and  L.  Packer.  1995.  Annual  variation 
in  survival  and  reproduction  of  the  primitively 
eusocial  sweat  bee  Halictus  ligatus  (Hymenoptera: 
Halictidae).  Canadian  Journal  of  Zoology  73: 
933-341. 

,  L.   Packer,  and  J.   Seger.   1995.   Unexpected 

patterns  of  parentage  and  relatedness  in  a  prim- 
itively eusocial  bee.  Nature  373:  239-241. 
-,  E.  J.  von  Wettberg,  and  A.  C.  Rutgers.  2003.  A 


novel  social  polymorphism  in  a  primitively  eu- 
social bee.  Proceedings  of  the  National  Academy  of 
Sciences  of  the  United  States  of  America  100: 
7175-7180'. 

Sakagami,  S.  F.  and  P.  A.  W.  Ebmer.  1979.  Halictus 
(Seladonia)  tumulorinu  higashi  ssp.  Nov.  from  the 
Northeastern  Palaearctic  (Hymenoptera:  Apoi- 
dea;  Halictidae).  Kontyu  47:  543-549. 

and  T.  Okazawa.  1985.  A  populous  nest  of  the 

Halictine  bee  Halictus  (Seladonia)  lutcsccus  from 
Guatemala  (Hymenoptera,  Halictidae).  Kontyu  53: 
645-651. 

Yanega,  D.  1988.  Social  plasticity  and  early-diapaus- 
ing  females  in  a  primitively  social  bee.  Proceedings 
of  the  National  Academy  of  Sciences  of  the  United 
States  of  America  85:  4374-4377. 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  293-296 

The  Status  of  Liris  magnificus  Kohl,  1884,  and  Trachogorytes  costaricae 
R.  Bohart,  2000  (Hymenoptera:  Crabronidae:  Crabroninae,  Bembicinae) 

WOJCIECH  J.  PULAWSKI 

Department  of  Entomology,  California  Academy  of  Sciences,  875  Howard  Street,  San  Francisco, 
California  94103,  USA;  email:  wpulawski@calacademy.org 


Abstract. — The  Australian  Liris  magnificus  Kohl,  1884,  currently  treated  as  a  subspecies  of  Liris 
haetnorrhoidalis  (Fabricius,  1803)  from  the  Palearctic  and  Afrotropical  Regions,  is  an  independent, 
full  species.  The  taxonomic  history  of  the  species  is  reviewed  and  the  differences  with 
haemorrhoidalis  are  discussed.  Trachogorytes  Bohart,  2000,  a  monotypic  genus  described  for 
Trachogorytes  costaricae  Bohart,  2000,  is  actually  a  junior  synonym  of  Mellinus  Fabricius,  1790. 
Mellimis  costaricae  Bohart,  2000,  comb,  nov.,  is  redescribed. 


Liris  magnificus  Kohl  by  Cardale  (1985)  and  Naumann  (1993). 

Liris  magnificus  Kohl,  1884:356,  E  (as  magnifica,  Bohart   and    Menke    (1976)'   on   the   other 

incorrect  original  termination).   Holotype  or  hand,  regarded   the  subspecific  status  of 

syntypes:   E,  northern  Australia:  no  specific  magnificus  as  tentative, 
locality  (Naturhistorisches  Museum  Wien).  -  Having  recently  examined  five  females 

Kohl,  1892:228  (in  key  to  world  Liris  s.s.);  Turner,  and  six  males  of  magnificus,  I  conclude  that 

1908:473  (as  new  synonym  of  Liris  haemorrhoi-  it  is  actually  a  full  species,  and  not  a  geo- 

dalis);   Dollfuss,    1989:10   (type   material   in  graphic  form  of  haemorrhoidalis.  The  differ- 

NHMW).  -  As  Larra  magnifica:  Kohl,  1885:245  u  .  .u  (  n  T 

;  ,  ,     ,,.       r        ,  ,  r       x  ences    between   them   are   as   follows,    in 

(new  combination,  m  checklist  of  world  Larra);  .r.  ,  ..  ...  ,    ,, 

^  ,,     T  1on„„„   ,.         .  ,         ,         .,  magnificus,   the   median   swelling   of   the 

Dalla   Torre,    1897:669   (in   catalog   of  world  *    J         '  & 

Hymenoptera).  -  As  Liris  haemorrhoidalis  Perioral  collar  is  wider  (Fig.  2b);  in  the 
magnifica:  Williams,  1928:49  (new  status,  nest-  female,    the    carina    emerging    from    the 
ing  habits);  Bohart  and  Menke,  1976:245  (as  clypeal  lobe  corner  is  about  twice  as  long 
tentative  subspecies  of  Liris  haemorrhoidalis);  as  the  midocellar  width  (Fig.  2a)  and  the 
Cardale,   1985:235   (in  catalog  of  Australian  dark  apical  coloration  of  fore  wing  does 
Sphecidae);  Naumann,   1993:185   (Australia:  not  extend  into  the  cell  area  (Fig.  2c);  the 
Queensland:  Heathlands  area  in  Cape  York).  male  hind  tarsomere  II  is  simple,  as  in  most 
Lin's  magnificus  was  described  as  a  full  of  the  congeners;  in  most  males  the  hind 
species,  but  was  synonymized  with  hue-  coxa   is   concave   ventrally   and   carinate 
morrhoidalis  (Fabricius)  by  Turner  (1908).  along  inner  margin  (Fig.  2d),  but  slightly 
Surprisingly,  the  author  stated  "I  cannot  convex  ventrally  and  obtuse  along  inner 
detect  any  appreciable  difference  in  the  margin   in   one    specimen    from   Wonga 
male",  but  in  fact  the  males  are  strikingly  Beach.  In  haemorrhoidalis,  the  median  swell- 
different  (see  below).  Both  species,  howev-  ing   of   the   pronotal    collar   is   narrower 
er,  are  similar  in  having  a  non-emarginate  (Fig-  lb);  in  the  female,  the  carina  emerging 
posterior  mandibular  margin,  red  legs,  and  horn  the  clypeal  lobe  corner  is  about  as 
conspicuously  golden  body  setae,  a  combi-  long  as  midocellar  width  (Fig.  la)  and  the 
nation  unique  within  the  genus.  Williams  dark   apical    area    of    fore    wing    covers 
(1928)  treated  magnificus  as  a  subspecies  of  marginal    as   well    as   second    and    third 
haemorrhoidalis,  an  interpretation  followed  submarginal  cells  (Fig.  lc);  in  the  male  the 


294 


Journal  of  Hymenoptera  Research 


'  -sF&.iiiiiiB  ■...■— ■■*•" 

^  i ^i\  ■■  - «&.' 

wFm' 

< 


■"I 


1  mm 


* 


1  mm 


Fig.  1.     Liris  haemorrhoidalis  (Fabricius):  a  -  lateral  carina  of  female  clypeal  lobe  in  oblique  view;  b  -  female 
pronotum;  c  -  apical  half  of  female  fore  wing;  d  -  male  hind  tarsomere  II. 


Fig.  :       Liris  magnificus  Turner:  a  -  lateral  carina  of  female  clypeal  lobe  in  oblique  view;  b  -  female  pronotum;  c 
-  apical  half  of  female  fore  wing;  d  -  male  hind  coxa. 


Volume  16,  Number  2,  2007 


295 


hind  coxa  is  neither  concave  nor  carinate, 
and  hind  tarsomere  II  is  conspicuously 
expanded  (Fig.  Id),  a  unique  such  feature 
in  the  genus.  The  genitalia  appear  identical 
in  both  species.  The  two  species  do  not 
occur  sympatrically:  Lin's  magnificus  is 
known  only  from  Australia,  whereas  hae- 
morrhoidalis  occurs  throughout  Africa, 
Spain,  the  Canary  Islands,  and  southwest- 
ern Asia  to  western  India  and  Sri  Lanka. 

Records  (all  specimens  are  at  the  California 
Academy  of  Sciences). — AUSTRALIA:  Queens- 
land: Armstrong  Beach  ca  15  km  E  Sarina  at 
21  27.3'S  149  17.5'E,  29  Oct.  2006,  W.J.  Pulawski  (1 
e?);  Balgal  Beach  51  km  NW  Townsville  at 
19  02.5'S  146°25.2'E,  18  May  2007,  V.E.  Ahrens 
and  W.J.  Pulawski  (1  9);  Blacks  Beach  ca  8  km  N 
Mackay  at  21  03.6'S  149  ll'E,  1  Dec  2006,  W.J. 
Pulawski  (1  J);  Burdekin  River  20  km  NE 
Charters  Towers  at  20  00.1 'S  146  26.3'E,  26  Nov 
2006,  W.J.  Pulawski,  and  21-22  May  2007,  V.E. 
Ahrens  and  W.J.  Pulawski  (1  9,  1  $);  Crystal 
Cascades  10  km  W  Cairns,  9-10  July  1983,  TW. 
Davies  (1  9);  69  road  km  WNW  Mount  Carbine  at 
16T3.2'S  144°43.8'E,  13  May  2007,  V.E.  Ahrens 
and  W.J.  Pulawski  (1  9);  Wonga  Beach  11  km  NNE 
Mossman  at  16  19.9'S  14525.3',  W.J.  Pulawski,  19 
Nov  2006  (1  9,  1  o)  and  21  Nov  2006  (1  S). 

Mellinus  costaricae  (R.  Bohart,  2000), 
new  combination 

Trachogon/tes  costaricae  R.  Bohart,  2000:168,  9. 
Holotype:  9/  Costa  Rica:  Puntarenas:  San  Vito 
(University  of  California,  Davis).  -  Amarante, 
2002:19  (in  catalog  of  Neotropical  Crabronidae). 

R.  Bohart  (2000)  published  an  important 
revision  of  the  Neotropical  Gorytini  in 
which  he  described  eight  new  genera  and 
a  number  of  new  species.  One  of  them  was 
Trachogorytes  costaricae,  based  on  a  single 
female  from  Costa  Rica.  I  have  examined 
that  specimen  and  found  that  it  is  a  member 
of  Mel  Hints  based  on  the  wing  venation 
(second  submarginal  cell  not  receiving  any 
of  the  recurrent  veins),  non-emarginate 
posterior  mandibular  margin,  short  tongue, 
absence  of  an  omalus  and  oblique  scutal 
carina,  raised  and  well  separated  pronotal 
collar,  scutellum,  and  metanotum,  evident 


Fig.  3.  Mellinus  costaricae  (Bohart),  holorvpe:  a  - 
whole  body  in  lateral  view;  b  -  mesopleuron;  c  - 
propodeal  enclosure. 

notaulus,  propodeal  dorsum  with  well-de- 
fined enclosure,  submarginal  cell  III  long 
and  distally  acute,  mid-coxa  simple,  pres- 
ence of  two  mid-tibial  spurs,  and  a  pedun- 
culate gaster  (Bohart  and  Menke  1976). 

In  Menke's  key  (1996)  to  Neotropical 
Mellinus,  this  species  runs  to  henseni  Menke. 
It  differs  from  henseni  and  all  other  currently 


296 


Journal  of  Hymenoptera  Research 


recognized  Mellinus  (Siri  and  Bohart  1974, 
Menke  1996)  by  its  unique  sculpture:  the 
mesopleuron  is  longitudinally  ridged  in  the 
posterior  half  (Fig.  3b)  rather  than  punctate 
or  uniformly  microsculprured,  the  propo- 
deal  enclosure  is  all  coarsely  rugose  (except 
at  the  very  apex),  the  propodeal  side  is 
longitudinally  ridged,  and  the  propodeal 
posterior  surface  is  rugose  (Fig.  3c),  not 
punctate,  as  stated  in  the  original  descrip- 
tion. Additionally,  the  tentorial  pit  is  closer 
to  the  antennal  socket  than  to  the  eye 
margin  and  the  propodeal  side  is  separated 
from  the  posterior  surface  by  a  conspicuous 
carina,  as  in  the  Palearctic  arvensis  (Lin- 
naeus) and  crabroneus  (Thunberg).  Most  of 
the  body  is  black  (Fig.  3a),  but  the  following 
are  whitish:  narrow  paraorbital  strip  in  the 
ventral  half  of  the  frons,  clypeus  (except 
along  frontoclypeal  margin),  scape  ventral- 
ly,  mandible  (except  apically),  mesally 
interrupted  fascia  on  pronotal  collar,  pro- 
notal  lobe  apically,  anterior  half  of  tegula, 
tiny  median  spot  on  metanotum,  tergum  I 
laterally  (except  in  basal  half)  and  apically, 
and  a  pair  of  preapical,  widely  separated 
spots  on  tergum  II.  The  femora  are  blackish 
basally,  then  reddish  brown  and  yellow;  the 
tibiae  are  reddish  brown  and  yellow;  and 
the  tarsi  are  yellow. 

ACKNOWLEDGMENTS 

I  thank  Mr.  Pavel  G.  Nemkov,  Vladivostok,  Russia, 
who  first  drew  my  attention  to  the  correct  generic 
position  of  Trachogorytes  costaricae  during  our  visit  to 
the  Bohart  Museum  of  Entomology,  University  of 
California,  Davis,  on  17  April  2007.  I  am  indebted  to 
Steven  L.  Heydon,  Bohart  Museum  of  Entomology, 
and  Brian  Harris,  United  States  National  Museum  of 
Natural  History,  for  lending  holotypes  of  Trachogor- 
ytes costaricae  and  Mellinus  henseni,  respectively. 
Robert  L.  Zuparko,  California  Academy  of  Sciences, 
and  Arnold  S.  Menke,  Bisbee,  Arizona,  kindly 
reviewed  a  draft  of  the  manuscript  and  made 
a  number  of  significant  improvements. 

LITERATURE  CITED 

Amarante,  S.  T.  P.  2002.  A  synonymic  catalog  of  the 
Neotropical  Crabronidae  and  Sphecidae  (Hyme- 
noptera: Apoidea).  Arquivos  de  Zoologia  37:  1-139. 


Bohart,  R.  M.  2000.  A  review  of  Gorytini  in  the 
Neotropical  Region  (Hymenoptera:  Sphecidae: 
Bembicinae).  Contributions  on  Entomology,  Interna- 
tional 4:  111-259. 

and  A.  S.  Menke.  1976.  Sphecid  Wasps  of  the 

World.  A  generic  revision.  University  of  California 
Press,  Berkeley,  Los  Angeles,  London.  1  color 
plate,  IX  +  695  pp. 

Cardale,  J.  1985.  Sphecidae.  Pp.  218-303  in  D.  W. 
Walton,  ed.  Zoological  Catalogue  of  Australia,  2. 
Hymenoptera.  Formicoidea,  Vespoidea  and  Sphecoidea. 
Australian  Government  Publishing  Service,  Can- 
berra, i-vi,  381  pp. 

Dalla  Torre,  C.  G.  1897.  Catalogus  Hymenopterorum 
hucusque  descriptorum  systematicus  et  synonymicus. 
Volumen  VIII:  Fossores  (Sphegidae).  Guilelmi 
Engelmann,  Lipsiae.  749  pp. 

Dollfuss,  H.  1989.  Verzeichnis  der  Grabwespentypen 
am  Naturhistorischen  Museum  in  Wien  (Hyme- 
noptera, Sphecidae).  Kataloge  der  wissenschaftlichen 
Sammlungen  des  Naturhistorischen  Museums  in 
Wien.  Entomologie  7  (4):  1-26. 

Kohl,  F.  F.  1884  (1883).  Neue  Hymenopteren  in  den 
Sammlungen  des  k.  k.  zool.  Hof-Cabinetes  zu 
Wien.  II.  Verhandlungen  der  kaiserlich-koniglichen 
Zoologisch-Botanischen  Gescllschaft  in  Wien  33: 
331-386,  pis.  XVIIa-XVIII. 

— .  1885  (1884).  Die  Gattungen  und  Arten  der 
Larriden  Autorum  [sic].  Verhandlungen  der  kaiser- 
lich-koniglichen Zoologisch-Botanischen  Gesellschaft 
in  Wien  34:  171-268,  pis.  VIII-IX,  327-454,  pis.  XI- 
XII. 

.  1892.  Neue  Hymenopterenformen.  Annalen  des 

k.k.  Naturhistorischen  Hofmuseums  7:  197-234,  pis. 
XIII-XV. 

Menke,  A.  S.  1996.  Neotropical  Mellinus:  a  review 
(Hymenoptera:  Sphecidae).  Memoirs  of  the 
Entomological  Society  of  Washington  17:  125- 
141. 

Naumann,  I.  D.  1993.  Results  for  aculeate  wasps.  Pp. 
175-187  in  I.  D.  Naumann,  E.  D.  Edwards,  T.  A. 
Weir,  and  D.  C.  F.  Rentz.  Insects  of  the  Heath- 
lands  area,  Cape  York  Peninsula,  Queensland.  Cape 
York  Peninsula  scientific  expedition.  Wet  Season 
1992.  Report.  The  Royal  Geographical  Society  of 
Queensland  Inc.  Vol.  2:  173-203. 

Turner,  R.  E.  1908.  Notes  on  the  Australian  fos- 
sorial  wasps  of  the  family  Sphegidae,  with 
descriptions  of  new  species.  Proceedings  of  the 
General  Meetings  for  Scientific  Business  of  the 
Zoological  Society  of  London  1908:  457-535,  pi. 
XXVI. 

Williams,  F.  X.  1928.  Studies  in  tropical  wasps  -  their 
hosts  and  associates  (with  descriptions  of  new 
species).  Bulletin.  Reports  of  Work  of  the  Experiment 
Station  of  the  Hawaiian  Sugar  Planters'  Association. 
Entomological  Series  19:  1-179. 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  297-310 

Interspecific  Variation  in  Hunting  Behavior  of  Pepsis  grossa  (Fabricius) 
and  Pepsis  thisbe  Lucas  (Hymenoptera:  Pompilidae):  A  field  study 

Fred  Punzo 

Department  of  Biology,  Box  5F,  University  of  Tampa,  401  W.  Kennedy  Blvd.,  Tampa, 

Florida  33606,  USA;  email:  fpunzo@ut.edu 


Abstract. — Field  studies  were  conducted  on  encounters  between  the  spider  wasps  Pepsis  grossa 
(Fabricius)  and  P.  thisbe  Lucas,  and  females  of  their  host  spider,  Aphonopelma  steindachneri 
(Ausserer)  (Theraphosidae),  in  Big  Bend  National  Park,  Texas.  Females  of  P.  grossa  were 
significantly  larger  than  those  of  P.  thisbe.  Number  of  eggs  found  in  ovarioles  of  P.  grossa  and  P. 
thisbe  ranged  from  6-14  and  3-12,  with  a  mean  of  11.3  and  8.4,  respectively.  Behavioral  acts 
comprising  hunting  behavior  of  both  species  included  antennation  of  a  spider's  burrow  (BA), 
evicting  spiders  from  their  burrow  (EVB),  initial  approach  and  antennation  of  spider  (AA),  moving 
away  and  grooming  (MG1),  attack  and  paralysis  (AP),  moving  away/grooming  (MG2),  drinking 
behavior  (DB),  burial  of  spider  and  egg  deposition  (BO),  and  closure  of  the  burrow  entrance  (BC). 
Antennae  of  most  wasps  made  initial  contact  with  the  forelegs  or  palps  of  a  spider.  During  AP, 
wasps  typically  grasped  leg  3  or  4  of  the  host  before  inserting  their  stings.  Most  wasps  of  P.  grossa 
(78%)  inserted  their  sting  into  the  intersegmental  membrane  between  the  sternum  and  coxa  2  of  the 
spider;  88%  of  P.  thisbe  females  chose  a  site  between  the  sternum  and  coxa  1.  Only  33  and  26%  of  P. 
grossa  and  P.  thisbe,  respectively,  drank  fluids  from  a  spider's  mouth  or  from  sting  insertion  site 
(LB).  Pepsis  thisbe  required  significantly  more  time  (mean:  129.1  min)  to  complete  all  behavioral  acts 
of  hunting  than  P.  grossa  (mean:  101.4  min).  Wasps  were  successful  in  paralyzing  spiders  in  all 
observed  encounters,  and  no  spider  attempted  to  attack  a  wasp. 


Aculeate  spider  wasps  of  the  genus 
Pepsis  (Hymenoptera:  Pompilidae)  include 
at  least  133  species  varying  in  size  from 
>60  mm  to  <12  mm  in  length  (Vardy 
2000).  Most  members  of  this  genus  have 
been  referred  to  as  tarantula  hawk  wasps 
because  females  selectively  hunt  mygalo- 
morph  spiders  of  the  family  Theraphosidae 
(Cazier  and  Mortenson  1964,  Punzo  and 
Garman  1989,  Vardy  2002).  They  paralyze 
and  store  these  spiders  (hosts)  in  under- 
ground nests  (Williams  1956,  Punzo  and 
Ludwig  2005)  as  a  food  source  for  their 
carnivorous  larvae  (Punzo  1994a).  They 
occur  throughout  the  New  World,  from 
the  United  States  and  West  Indies,  south  to 
Patagonia  (Hurd  1952,  Vardy  2000). 

Pepsis  grossa  (Fabricius  1798)  and  P.  thisbe 
Lucas  (1895)  are  large,  long-legged  wasps 
and  are  conspicuous  components  of  the 


arthropod  fauna  of  desert  regions  in  the 
southwestern  United  States  and  northern 
Mexico  (Hurd  1952,  Vardy  2000,  Punzo, 
1994b,  2006a).  In  Big  Bend  National  Park 
(BBNP;  Brewster  County,  Texas,  USA), 
females  of  these  two  species,  as  well  as  P. 
mildei  Stal  (1844)  selectively  hunt  and 
paralyze  the  large  theraphosid  spider, 
Aphonopelma  steindachneri  (Ausserer  1929) 
which  they  use  as  a  host  for  their  de- 
veloping larvae  (Punzo  2005a). 

Adult  wasps  feed  on  nectar  which  is 
obtained  from  flowers  of  a  variety  of  plants 
(Evans  and  West-Eberhard  1970,  Punzo 
2000,  2006a).  In  BBNP,  where  ambient 
temperatures  in  late  spring  and  summer 
may  exceed  43  C,  it  is  not  uncommon  for 
females  to  fly  over  considerable  distances 
during  daylight  hours  in  search  for  flowers 
and  host  spiders  (Punzo  1994b,  Schmidt 


298 


Journal  of  Hymenoptera  Research 


2004).  It  is  important  for  females  to  obtain 
adequate  amounts  of  required  nutrients 
because  insect  flight  places  high  metabolic 
demands  on  insects  (Nation  2002). 

Females  of  Aphonopelma  steindachneri 
(Ausserer),  like  other  spiders  of  this  genus, 
excavate  burrows  (or  occupy  abandoned 
rodent  burrows)  where  they  remain  for 
most  of  their  lives  (Baerg  1958,  Gabel  1972, 
Punzo  2007a).  Females  seize  prey  that 
approaches  close  to  the  burrow  entrance. 
Adult  males  actively  wander  over  the 
ground  surface  during  the  mating  season 
when  they  search  for  conspecific  females 
(Punzo  2000,  2007b). 

Female  Pepsis  wasps  initiate  their  search 
for  suitable  spider  hosts  after  mating 
(Punzo  1994b,  2006a,b).  They  typically  fly 
over  the  ground  and  are  thought  to  detect 
occupied  spider  burrows  at  a  distance 
using  visual  and/or  odor  cues.  They  in- 
termittently interrupt  flight  by  landing  on 
the  ground  and  walk  rapidly  over  the 
surface,  tapping  the  ground  surface  fre- 
quently with  their  antennae  as  they  search 
for  spider  burrows  (Kurczewski  and  Kurc- 
zewski  1968).  When  a  suitable  spider 
burrow  is  located,  the  female  wasp  typi- 
cally stops  at  the  entrance  and  taps  the 
edge  of  the  opening  with  her  antennae 
(burrow  antenna tion,  BA).  After  a  variable 
period  of  time,  she  cuts  through  the  silk 
covering  over  the  burrow  entrance  with 
her  mandibles,  enters  the  burrow,  and 
usually  forces  the  spider  out  of  its  burrow 
and  onto  the  surface  (EVB,  eviction  behav- 
ior). Male  theraphosids  are  usually  en- 
countered as  they  move  about  searching 
for  food  and  mates  (Minch  1979,  Punzo 
2005b,  2007b).  Unlike  burrow-dwelling 
females,  males  of  the  genus  Aphonopelma 
from  dessert  regions  usually  seek  shelter 
within  or  under  rock  crevices,  or  under 
surface  debris  (Smith  1994,  Punzo  and 
Henderson  1999). 

A  specific  sequence  of  behavioral  acts 
are  exhibited  once  a  spider  has  been  forced 
to  the  ground  surface.  These  acts  comprise 
the  overall  attack  behavior  of  Pepsis  wasps 


found  in  the  tropics  and  desert  regions  of 
the  southwestern  United  States  (Petrunke- 
vitch  1926,  1952,  Cazier  and  Mortenson 
1964,  Punzo  and  Garman  1989,  Punzo  1991, 
1994b,  2005a,c).  A  female  wasp  typically 
approaches  the  spider  and  touches  its  body 
surface  with  her  antennae  (approach  and 
antennation,  AA)  (Punzo  and  Garman 
1989).  In  some  cases,  the  spider  does  not 
move  away,  although  it  may  twitch  one  of 
its  forelegs  or  raise  its  palps  off  the  ground 
(Punzo  2007b).  In  other  instances,  tactile 
stimulation  by  a  wasp  elicits  a  threat 
posture  from  the  spider  which  elevates 
the  anterior  legs  and  exposes  its  fangs 
(Petrunkevitch  1952,  Punzo  1994b).  How- 
ever, the  spider  rarely  strikes  at  the  wasp, 
although  if  presented  with  another  insect 
(cricket,  etc.)  it  typically  strikes  quickly, 
seizing  the  insect  and  eating  it.  It  has  been 
suggested  that  chemosensory  cues  associ- 
ated with  the  wasp's  cuticle  inhibit  the 
spider's  strike  response  (Punzo  2000). 

Once  a  spider  has  been  identified  as 
a  suitable  host,  the  wasp  moves  a  short 
distance  away  and  exhibits  grooming 
behavior  by  passing  its  antennae  through 
the  mandibles.  This  behavioral  component 
is  known  as  'moving  away  and  grooming' 
(MG1,  Punzo  1991).  After  a  few  minutes, 
the  wasp  turns  to  face  its  host  and  then 
walks  under  the  ventral  body  region  of  the 
spider.  In  response  to  this,  spiders  usually 
extend  their  legs,  elevating  their  body  off 
the  ground.  The  wasp  then  grasps  one  of 
the  spider's  legs  and  quickly  inserts  its 
sting  through  the  ventral  body  region  into 
the  prosomal  nerve  mass  resulting  in 
a  rapid  paralysis  of  the  spider.  This 
component  of  hunting  is  referred  to  as 
attack  and  paralysis  (AP).  Once  a  spider  is 
immobilized,  the  wasp  moves  away  and 
repeats  the  grooming  sequence  described 
above  (MG2).  It  then  returns  to  the  spider 
and  in  some  instances  may  either  drink 
fluids  from  the  spider's  mouth  cavity  or 
drink  spider  hemolymph  that  leaks  out  of 
the  puncture  wound  made  by  the  insertion 
of  the  sting.  This  is  known  as  drinking  or 


Volume  16,  Number  2,  2007  299 

lapping  behavior,  DB  (Punzo  2000).  How-  air   temperatures   ranging   from   5.4  C   in 

ever,  because  DB  does  not  occur  in  most  January  to  33.8  C  in  August  (US  Dept.  of 

encounters,  the  question  arises  as  to  what  Interior  2005).  Annual  rainfall  is  between 

factor(s)  may  be  responsible  for  its  occur-  13.8-30.1  cm,  depending  on  location  and 

rence.  altitude,  with  65-70%  occurring  from  May 

The  paralyzed  host  is  then  dragged  into  through  October  (Parent  1996).  Topogra- 

the  spider's  burrow  (or  one  excavated  by  phy  of  the  Park  is  diverse  and  includes 

the  wasp)  and  a  single  egg  is  deposited  on  gypsum   formations,   igneous   rocks,   and 

the  ventral  surface  of  the  spider's  abdomen  limestone  deposits  that  provide  different 

(burial   and   oviposition,   BO).   The   wasp  substrates  including  alluvial  fans,  gypsum 

then   closes    the   burrow    entrance    (BC,  flats,  saline  playas,  siliceous  and  gypsum 

burrow  closure)  using  soil  particles  and  dunes,    fine-textured    basins,    canyons, 

small  pebbles  and  flies  off  to  search  for  mountain  ridges,  and  freshwater  springs 

another  host.  and  seeps,  all  supporting  a  diverse  plant 

Although  behavioral  acts  of  the  hunting  fauna  categorized  within  distinctive  vege- 

sequence  have  been  studied,  most  observa-  tative  zones  (Powell  1988). 
tions  are  based  on  laboratory  encounters 

between   wasps   and   hosts   (Punzo   1991,  MATERIALS  AND  METHODS 

1994b,  2007).  Few  detailed  observations  of  I  conducted  field  studies  over  a  4-year 

encounters   in    the   field   have   been   de-  period  (2002-2005)  within  BBNP  from  May 

scribed.  In  addition,  there  are  a  number  through   September,   when   male   wasps 

of   questions   that   remain.   For   example,  were  establishing  perch  sites  (territories) 

when   attacking   a    spider   does   a   wasp  and  females  were  searching  for  spiders, 

exhibit  any  preference  for  grasping  a  par-  Adults    of   P.   grossa    and    P.thisbe   were 

ticular  leg?  Are  there  specific  sites  on  the  observed  within  a  5-km  radius  of  Tornillo 

spider's  body  where   a  wasp   inserts   its  Flat  (TF;  29  01'N,  102  59W),  a  site  where 

sting?  How  commonly  does  lapping  be-  both  of  these  species  are  abundant,  as  well 

havior  occur?  Is  there  a  preference  for  the  as  their  host  spider,  A.  steindachneri,  are 

site  at  which  it  occurs?  Finally,  are  there  abundant    (Punzo    2000,    2007b).    I    had 

interspecific   differences   associated    with  located  and  marked  numerous  occupied 

these  behaviors?  The  present  study  was  tarantula  burrows  during  previous  field 

conducted    in   order   to   analyze   hunting  studies  in  this  area  over  the  last  12  years  (;/ 

behavior  of  P.  grossa  and  P.  thisbe  under  =  946).  Females  of  A.  steindachneri  typically 

natural  conditions  and   to  address  these  remain  within  a  single  burrow  for  most  of 

questions  in  these  two  sympatric  species  of  their  adult  lives  (Punzo,  unpubl.  data).  As 

wasps  that  are  found  in  similar  microhab-  a    result,    I   knew    the    locations   of   host 

itats  in  Big  Bend  National  Park  (BBNP),  spiders  and  concentrated  my  field  observa- 

where  both  utilize  A.  steindachneri  as  a  host,  tions  at  these  burrow  sites.  Voucher  speci- 
mens of  wasps,  wasp  eggs,  and  spiders 

DESCRIPTION  OF  GENERAL  have  been  deposited   in  the  invertebrate 

STUDY  AREA  collection  at  BBNP. 

Pepsis  grossa  and  P.  thisbe  occur  through-  Based  on  my  knowledge  of  locations  for 

out   Big   Bend   National   Park   (BBNP)    is  burrows  occupied  by  a  female  tarantula,  I 

located  in  Brewster  County,  Texas,  and  lies  examined   148  burrows  whose  entrances 

within  the  northern  region  of  the  Chihua-  had  been  closed   from   previously  unob- 

huan   Desert.    Its   western,   southern   and  served  encounters  with   wasps  to  deter- 

eastern  boundaries  are  bordered  by  the  Rio  mine   whether   it   contained   a   paralyzed 

Grande  River.  Climatic  conditions  range  host.  During  the  course  of  this  study  I  also 

from  arid  to  semiarid,  with  mean  monthly  monitored  96  burrows  containing  an  adult 


300  Journal  of  Hymenoptera  Research 

female  spider,  and  observed  54  and  42  ses.   Another  Pepsis  wasp,  P.   mildei  also 

encounters  between  a  wasp  and  spider  for  occurs   at   TF,   but   is   far   less   abundant 

P.  grossa  and  P.  thisbe,  respectively.  Because  (Punzo,  unpubl.  data).  After  measurements 

encounters  between  a  wasp  and  a  male  were  recorded,  each  wasp  was  preserved 

spider   usually   occur   while   the   male   is  in  70%  ethanol  for  subsequent  determina- 

wandering  over  the  ground  surface,  op-  tion  of  number  of  eggs  in  ovarioles.  Re- 

portunities   to   witness   such   encounters  moval   of  wasps  ensured   that  the  same 

occur   far   less   frequent.   Therefore,   only  wasp  was  not  involved  in  more  than  one 

encounters  involving  a  female  spider  were  encounter  for  the  data  set. 

used  for  analyses.  For  each  encounter  I  observed  the  general 

I  observed  all  encounters  at  close  range  behavior  of  the  wasp  as  it  approached  the 

(1-2  m  from  combatants),  and  used  a  35-  burrow  entrance  and  interacted  with  a  spi- 

mm    Nikon   FE2   camera    to   photograph  der,  as  well  as  the  concomitant  behaviors 

some  of  the  encounters.  Encounters  were  exhibited  by  the  host.  I  recorded:  (1)  the 

observed    during   daylight   and    evening  amount  of  time  required  to  complete  the 

hours    (0830-0200  h,    Central    Standard  hunting  sequence  (from  initial  contact  to 

Time).    At    the   end    of   each    encounter  closure   of  the   burrow),   as   well   as   all 

(following  burial,  oviposition,  and  closure  behavioral  components  of  hunting,  using 

of  the  burrow)  I  collected  the  wasp  with  a   stopwatch;   (2)   which   spider   leg   was 

a   sweep   net   and    anesthetized    it   using  initially  seized  by  the  wasp  as  it  attempted 

a  portable  C02  cartridge.  Once  inactivated,  to  insert  its  sting;  (3)  site  at  which  wasp 

I  verified  species  identification  and  used  sting  had  been  inserted  into  the  body  of  the 

a  Unitron  dissecting  microscope  fitted  with  spider;   and   (4)   whether   or  not  lapping 

an    ocular    micrometer    to    record    body  behavior  occurred,  and  if  so,  where, 

length  and  head  capsule  width.  Although  All  statistical  procedures  followed  those 

Aphonopelma  steindachneri  is  the  only  ther-  described  by  Sokal  and  Rohlf  (1995).  All 

aphosid   known  to  occur  at  the  TF  site  data  conformed  to  conditions  of  normality 

(Smith  1994,  Punzo  2007b),  I  opened  each  as   assessed    using   a    Bartlett's    test   for 

burrow  to  verify  species  identification  of  homogeneity  of  variances  and  a  G-test  for 

paralyzed  spiders.  I  examined  the  body  normality.  Comparisons  on  means  for  mor- 

surface  of  paralyzed  spiders  with  a  dissect-  phometric  data  between  the  two  species  of 

ing  microscope  in  order  to  locate  the  site  wasps   as   well   as   for  male   and   female 

where  the  wasp's  sting  had  been  inserted  spiders  were  tested  using  a  t  test.  Differ- 

into    the   host   during    the   paralyzation  ences  between  the  proportion  of  burrows 

sequence.  I  also  recorded  the  width  of  the  containing  paralyzed  males  versus  females, 

carapace   and    total   body   length   to   the  and  frequency  at  which  specific  spider  legs 

nearest   0.1mm   using   a    digital   caliper,  were  grasped  by  a  wasp  during  initiation  of 

After  removing  the  wasp's  egg,  I  recorded  attack  behavior,  were  tested  using  a  Chi 

weight    of   wasp    eggs    (to    the    nearest  Square  test  (X2).  Comparisons  between  time 

0.01  mg)    and    spiders    (to    the    nearest  required  by  each  wasp  species  to  complete 

0.01  g)  using  a  portable  electronic  balance,  overall  hunting  sequence  was  tested  using 

Egg  length  and  width  were  recorded  to  the  an   analysis   of  variance   (ANOVA),   and 

nearest  0.01  mm  using  a  dissecting  micro-  a   Scheffe   F   test   was   used   for   ad   hoc 

scope.  comparisons  between  individual  behavioral 

The  site  of  the  puncture  wound  could  be  components  of  hunting, 
readily    identified    by    hemolymph    that 

oozed    out    of    the    wound.    Only    data  RESULTS 

obtained  for  P.  grossa,  P.   thisbe  and  Of  the  148  burrows  whose  entrances  had 

host  A.  steindachneri  were  used  for  anal}  been  closed  with  soil  (following  previously 


Volume  16,  Number  2,  2007 


301 


Table  1.  Morphometric  data  on  females  of  Pepsis  grossa  (n  =  54)  and  P.  thisbe  (n  =  42)  and  female  host 
spiders,  Apkonopelma  steindachneri  (n  =  96  females,  46  males)  at  Tornillo  Flat,  Big  Bend  National  Park,  Texas. 
Data  from  wasps  and  spiders  examined  from  2002  to  2005.  Data  expressed  as  means;  values  in  parentheses 
represent  (±SE).  BL  (body  length);  HCW  (head  capsule  width);  CW  (cephalothorax  width);  BW  (body  weight); 
NEO  (number  of  eggs  found  in  ovarioles).  Values  in  rows  followed  by  a  different  letter  are  statistically 
significant  (t  tests;  P  <  0.05). 


Pepsis  grossa 

P.  thisbe 

A. 

steindachneri 

BL  (mm) 
HCW  (mm) 
NEO 

CW  (mm) 
BW(g) 

42.7a  (2.4) 

4.9a  (0.2) 

11.3a  (2.4) 

35.9b  (1.7) 
3.9b  (0.3) 
8.4b  (0.7) 

Males 

13.4a  (0.7) 
6.8a  (0.5) 

Females 

15.2b  (0.5) 
10.8b  (1.1) 

unobserved  encounters),  128  (87%)  con- 
tained a  paralyzed  spider.  Although  the 
species  of  wasp  responsible  for  the  paral- 
ysis cannot  be  known  unless  the  offspring 
is  reared,  87  of  these  128  burrows  (68%) 
contained  a  female  spider,  and  32%  held 
a  male  (Chi  Square  test:  X2  =  10.89,  P  < 
0.03).  Paralyzed  male  spiders  ranged  in 
weight  from  5.2-6.4  g  (mean:  5.77  ±  0.42), 
while  the  range  was  6.9-10.4  g  (mean:  7.94 
±  0.37)  for  females.  For  data  obtained  from 
observed  encounters,  female  spiders  para- 
lyzed by  P.  grossa  and  P.  thisbe  had  a  mean 
weight  of  8.7  ±  0.83  g  (range:  6.2-10.6)  and 
7.7  ±  0.64  g  (range:  5.7-10.7),  respectively. 
Mean  values  for  length,  width,  and  weight 
for  eggs  of  P.  grossa  were  4.34  ±  0.03  mm, 
1.34  ±  0.01  mm,  and  7.14  ±  0.31  mg,  as 
compared  to  4.29  ±  0.05  mm,  1.24  ± 
0.02  mm,  and  6.88  ±  0.41  mg  for  P.  thisbe. 

Morphometric  data  for  wasps  observed 
encountering  spiders,  as  well  as  for  hosts, 
are  shown  in  Table  1.  Concerning  host 
spiders,  based  on  width  of  cephalothorax 
(t  =  3.04,  P  <  0.05)  and  body  weight  (t  = 
2.46,  P  <  0.05),  females  were  significantly 
larger  than  males.  For  the  two  species  of 
wasps,  females  of  P.  grossa  were  signifi- 
cantly larger  than  those  of  P.  thisbe,  based 
on  body  length  (t  =  3.35,  P  <  0.05)  and 
head  capsule  width  (t  =  2.88,  P  <  0.05). 

Data  on  wasps  collected  from  the  field 
indicated  that  the  mean  number  of  eggs 
found  in  ovarioles  for  P.  grossa  and  P.  thisbe 
was  11.3  and  8.4,  respectively  (Table  1), 


with  a  range  of  6-14  and  3-12.  Because 
wasps  require  a  host  for  each  egg,  and 
number  of  previous  encounters  with  a  spi- 
der was  unknown,  the  number  of  eggs 
produced  by  each  species  of  wasp  prior  to 
any  hunting  experience  could  not  be  de- 
termined from  field  data.  However,  I  have 
reared  both  species  of  wasps  from  larvae 
feeding  on  A.  steindachneri  females  (mean 
weight:  9.89  ±  0.61  g)  in  the  laboratory  and 
found  that  non-mated  P.  grossa  (10-12  days 
of  age)  produced  9-21  eggs /female  (mean: 
16.2  +  3.2  g  SE,  n  =  78)  as  compared  to  4-14 
(mean:  10.2  +  2.2,  n  =  38)  for  P.  thisbe  (t  = 
7.09,  P  <  0.05)  (Punzo,  unpubl.  data). 

For  all  observed  encounters  between  P. 
grossa  or  P.  thisbe  and  a  host  spider,  wasps 
entered  an  occupied  burrow  by  cutting 
through  the  silk  covering  over  the  burrow 
entrance  and  then  forced  the  host  to  the 
surface.  Attack  and  paralysis  never  oc- 
curred within  the  burrow.  A  wasp  would 
typically  approach  the  entrance  of  a  spider 
burrow  and  tap  its  antennae  along  the  edge 
of  the  opening.  The  time  allocated  by  these 
wasps  for  each  behavioral  component  of 
hunting  behavior  is  shown  in  Table  2. 

Pepsis  thisbe  females  required  signifi- 
cantly more  time  to  complete  the  overall 
hunting  sequence  as  compared  to  P.  grossa 
(F  =  19.27,  P  <  0.05)  (Table  2).  No 
significant  interspecific  differences  were 
found  for  eviction  behavior  (EVB;  Scheffe 
F,  P  >  0.50)  or  moving  away  and  grooming 
(MG1,  MG2,  P  >  0.60).  Interspecific  differ- 


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Journal  of  Hymenoptera  Research 


Table  2.  Time  (in  min)  allocated  by  females  of 
Pepsis  grossa  (n  =  54)  and  P.  thisbe  («  =  42)  for  various 
behavioral  components  of  the  overall  hunting 
sequence  during  encounters  with  a  host  spider, 
Aphonopelma  steindachneri.  Data  are  expressed  as 
means;  numbers  in  parentheses  represent  tSE. 
Values  in  rows  followed  by  a  different  letter  are 
statistically  significant  (P  <  0.05).  BA  (burrow 
antennation);  EVB  (eviction  behavior);  AA  (approach 
and  antennation);  MG1,  MG2  (moving  away  and 
grooming);  AP  (antennation  and  paralysis);  LB 
(lapping  behavior);  BO  (burial  and  oviposition);  BC 
(burrow  closure). 


Behavioral  component 

Pepsis  grossa 

Pepsi's  thisbe 

BA 

4.8a  (0.8) 

7.6b  (1.1) 

EVB 

3.4a  (0.3) 

3.9a  (0.4) 

AA 

8.3a  (1.7) 

5.1b  (0.8) 

MG1 

4.6a  (0.4) 

5.2a  (0.3) 

AP 

1.8a  (0.2) 

2.3a  (0.5) 

MG2 

3.2a  (0.6) 

2.9a  (0.3) 

LB1 

3.6a  (1.1) 

5.1b  (1.8) 

BO2 

14.4a  (3.5) 

23.2b  (4.6) 

BC 

57.3a  (7.1) 

73.8b  (6.9) 

Total: 

101.4a 

129.1b 

1  Lapping  behavior  occurred  in  18  of  54  encounters  for 
P.  grossa  (33.3%),  and  in  11  of  42  encounters  for  P. 
thisbe  (26.2%). 

2  Represents  data  for  situations  in  which  a  spider  was 
buried  in  its  own  burrow. 


ences  for  all  other  behavioral  components 
were  significant.  As  compared  to  P.  grossa, 
P.  thisbe  females  allocated  significantly- 
more  time  for  burrow  antennation  (BA; 
Scheffe  F  =  6.2,  P  <  0.05),  drinking 
behavior  (DB;  F  =  4.9,  P  <  0.05),  burial 
and  oviposition  (BO;  F  =  7.1,  P  <  0.05),  and 
burrow  closure  (BC;  F  =  6.9,  P  <  0.05),  and 
significantly  less  time  for  approach  and 
antennation  (AA;  F  =  5.5,  P  <  0.05). 

Wasps  of  both  species  approached  spi- 
ders that  they  had  forced  out  of  their 
burrows  and  then  tapped  the  spider's  body 
surface  with  their  antennae.  Antennae  of  P. 
grossa  and  P.  thisbe  initially  made  contact 
with  the  tarsus  of  one  of  the  spider's 
forelegs  in  50  of  54  (92.5%)  and  39  of  42 
(92.8%)  encounters,  respectively.  In  other 
cases,  the  antennae  initially  made  contact 
with  one  of  the  spider'spalps.  Subsequent- 
ly,   wasps    of   both    species    used    their 


antennae  to  explore  the  lateral  region  of 
a   spider's   cephalothorax   and   abdomen. 
During  antennation  by  P.  grossa,  34  of  54 
spiders  (63%)  exhibited  no  bodily  move- 
ments   as    compared    to    similar   values 
observed  for  P.  thisbe  (28  of  42,  67%,  P  > 
0.60).  For  encounters  with  P.  grossa,  other 
spiders   either   remained    stationary   but 
exhibited  slight  movements  of  their  foreleg 
(n  =  2,  4%)  or  an  elevation  of  the  palps 
(n  =  6,  11%),  while  the  remainder  (n  =  12, 
22%)  exhibited  a  threat  posture.  Compara- 
ble values  for  similar  responses  of  spiders 
to  P.  thisbe  were  5%  (n  =  2),  9%  (n  =  4),  and 
19%  (n  =  8),  respectively.  In  no  case  did 
a    spider   attempt   to   flee   back   into   its 
burrow   or  attack  the   wasp,   and   wasps 
'won'  all  observed  encounters. 

When  initiating  attack,  wasps  of  both 
species  showed  a  preference  for  grasping 
leg  3  or  4  of  the  spider  (Table  3).  Eight-one 
and  57%  of  P.  grossa  and  P.  thisbe,  re- 
spectively, exhibited  a  rapid  dash  under 
the  ventral  region  of  the  spider  and  used 
their  mandibles  to  grasp  leg  3  or  4  before 
attempting  to  insert  their  sting.  Leg  1  was 
never  grasped,  and  leg  2  in  only  2.4-7.4% 
of  encounters. 

Sting  insertion  sites  for  P.grossa  and  P. 
thisbe  are  shown  in  Fig.  1.  Examination  of 
spiders  post-paralysis  showed  that  78%  of 
P.  grossa  females  inserted  their  sting  into 
the  intersegmental  membrane  between  the 
sternum  and  coxa  2  of  the  spider,  and  22% 
between  the  sternum  and  pedipalp.  In 
comparison,  88  and  12%  of  P.  thisbe 
females,  respectively,  inserted  their  sting 
into  the  membrane  between  the  sternum 
and  coxa  1  or  at  the  junction  between  the 
abdomen  and  cephalothorax.  Mean  time 
that  elapsed  between  insertion  of  sting  and 
immobilization  (paralysis)  of  spider  (in- 
dicated by  curling  of  the  legs  under  the 
spider's  body)  was  6.2  s  ±  0.4  SE  (range:  4- 
8  s)  for  P.  grossa  and  12.8  s  ±  1.1  SE  (range: 
8-16  s)  for  P.  thisbe.  Regardless  of  insertion 
site,  there  was  no  significant  difference  in 
time  required  for  paralysis  for  either 
species  of  wasp. 


Volume  16,  Number  2,  2007 


303 


Table  3.  Leg  of  spider  (Aphonopelma  steindachneri) 
grasped  by  females  of  Pepsis  grossa  and  Pepsis  thisbe 
when  initiating  attack.  Data  derived  from  a  single 
observation  of  each  wasp/spider  encounter  for  54  and 
42  encounters,  respectively,  between  P.  grossa  and  P. 
thisbe,  and  a  host  spider.  Spider  legs  on  right  and  left 
side  of  the  body  (based  on  position  of  spider  when 
a  wasp  was  facing  it)  are  designated  as  R  and  L, 
respectively,  and  legs  are  numbered  1  (forelegs) 
through  4  (hindlegs).  Values  in  parentheses 
represent  frequency  of  occurrence  (%). 


Spider  leg  grasped 

Pepsis  grossa  (n  =  54) 

Pepsis  thisbe  (n  =  42) 

Rl 

0 

0 

LI 

0 

0 

R2 

4  (7.4) 

1  (2.4) 

L2 

0 

1  (2.4) 

R3 

20  (37) 

14  (33.3) 

L3 

24  (44.4) 

10  (23.8) 

R4 

4  (7.4) 

7  (16.6) 

L4 

2  (3.7) 

9  (21.4) 

Only  33.3  and  26.2%  of  P.  grossa  and  P. 
thisbe  females,  respectively,  exhibited 
drinking  behavior  (DB,  Table  2).  Eleven  of 
18  (61%)  females  of  P.  grossa  were  observed 
to  drink  fluid  oozing  from  the  wound  site 
(sting  insertion  site),  while  7  (39%)  wasps 
drank  fluids  from  the  spider's  mouth 
cavity.  For  P. thisbe,  the  percentage  of 
females  that  engaged  in  DB  was  8  of  11 
(73%)  and  3  of  11  (27%)  for  the  wound  site 
and  mouth  cavity,  respectively. 

DISCUSSION 

Physical  dimensions  and  number  of  eggs 
for  P.  grossa  and  P.  thisbe  were  similar  to 
values  reported  for  other  species  of  Pepsis 
wasps  of  similar  size.  For  example,  eggs  of 
P.  cerberus  Lucas  and  P.  mexicana  Lucas 
from  another  area  of  BBNP  ranged  from 
4.19-4.29  and  4.25  vs.  4.31  mm  (length), 
1.26-1.38  and  1.18-1.29  mm  (width),  and 
7.18-7.32  and  7.06-7.17  g  (weight),  respec- 
tively (Punzo  2005c).  In  like  manner, 
number  of  eggs  produced  per  female  for 
P.  grossa  and  P. thisbe  were  similar  to  values 
reported  for  other  Pepsis  wasps  which 
range  from  2-44/female  (Haupt  1952, 
Evans  and  West-Eberhard  1970,  Punzo 
2000,  2005c).  It  has  been  shown  that  the 


number  of  eggs  produced  by  Pepsis  females 
(Evans  1953,  Punzo  2005c),  as  well  as  in 
many  other  insects  (Price  1975,  Ito  1980, 
Nation  2002)  is  positively  correlated  with 
body  size. 

Although  a  majority  of  paralyzed  spi- 
ders found  with  an  attached  Pepsis  egg 
(where  encounters  with  a  wasp  had  not 
been  observed)  contained  female  spiders 
(68%),  almost  1/3  contained  a  male.  Thus, 
it  appears  that  Pepsis  wasps  are  opportu- 
nistic hunters  and  will  readily  attack  a  male 
tarantula  even  though  males  are  usually 
smaller  than  females.  This  is  in  general 
agreement  with  previous  laboratory  or 
field  observations  showing  that  Pepsis 
wasps  will  attack,  paralyze,  and  deposit 
an  egg  on  male  and  female  theraphosid 
hosts  (Kurczewski  and  Kurczewski  1968, 
Punzo  2000).  An  experimental  protocol 
that  might  allow  us  to  determine  whether 
female  Pepsis  wasps  have  any  'preference' 
for  spiders  of  different  sexes  would  be  to 
observe  the  response  of  mated  female 
wasps  when  given  a  choice  between  a  male 
and  female  spider.  If  naive  females  are 
used  (no  previous  encounter  with  a  spider), 
one  can  also  assess  whether  such  a  prefer- 
ence, if  exhibited,  had  an  innate  compo- 
nent. 

Little  information  is  available  on  the 
number  of  eggs  produced  per  female  for 
Pepsis  wasps.  A  previous  study  yielded 
some  data  for  pepsine  wasps  collected 
from  Persimmon  Gap,  a  site  48  km  to  the 
northeast  of  TF  that  also  lies  within  BBNP 
(Punzo  2005c).  Number  of  eggs  in  ovarioles 
from  wasps  collected  immediately  after 
mating  (before  they  started  to  hunt  for 
hosts)  ranged  from  5-26  (mean:  12.7  ±  2.8 
SE)  and  4-20  (mean:  11.8  ±  1.7)  eggs  per 
female,  for  P.  cerberus  and  P.  mexicana, 
respectively.  It  is  difficult  to  extrapolate 
and  compare  these  data  with  values 
reported  for  P.  grossa  (11.3)  and  P.  thisbe 
(8.4)  in  the  present  study  because  there  was 
no  way  of  knowing  how  many  prior 
encounters  these  wasps  had  with  a  host 
before  they  were  collected.  These  compare 


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Journal  of  Hymenoptera  Research 


Pedipalp 


Chelicera         Pedipalp 


Spinnerets 

Fig.  1.  Ventral  body  region  of  Aphonopelma  steindachneri  showing  insertion  sites  (wound  sites)  for  the  sting  of 
Pepsi's  grossa  and  P.  thisbe.  Solid  circle  and  square  (intersegmental  membrane  between  sternum  and  pedipalp, 
and  sternum  and  coxa  2,  respectively)  are  insertion  sites  for  P.  grossa:  solid  triangle  and  diamond  (membrane 
between  sternum  and  coxa  1,  and  at  junction  of  cephalothorax  and  abdomen,  respectively)  are  insertion  sites  for 
P.  thisbe.  Legs:  L1-L4. 


to  9-21  eggs  for  P.  grossa  and  4-14  for  P. 
thisbe  reared  in  the  laboratory  (Punzo, 
unpubl.  data). 

Spiders  paralyzed  by  P.  grossa  and  P. 
thisbe  varied  in  body  weight.  Previous 
research  has  demonstrated  that  adult  size 
(as  assessed  by  head  capsule  width,  length 
of  legs  or  wings)  attained  by  other  species 
of  spider  wasps  is  positively  correlated 
with  the  mass  attained  by  their  last-instar 
larvae  (Vinson  1984,  Punzo  2005c),  which 
in  turn  is  positively  correlated  with  the 


mass  of  the  spider  that  the  larvae  fed  on 
(Price  1997,  Punzo  2005c).  This  most  likely 
accounts  for  the  ranges  in  adult  size 
observed  in  male  and  female  Pepisis  wasps 
in  the  field.  Although  empirical  evidence  is 
lacking,  it  would  be  interesting  to  de- 
termine to  what  extent  (if  any)  a  decision 
made  by  a  female  Pepsis  wasp  concerning 
size  of  host  suitable  for  attack  may  be 
influenced  by  the  wasp's  size. 

There  are  relatively  few  detailed  obser- 
vations and  analyses  on  foraging  (Cazier 


Volume  16,  Number  2,  2007 


305 


and  Mortenson  1964,  Punzo  and  Ludwig 
2005,  Punzo  2006a,b),  territoriality  (Rau 
and  Rau  1918,  Punzo  2000),  dispersal 
(Evans  and  West-Eberhard  1970),  diel 
periodicity  (Punzo  2005c),  and  hunting 
behavior  (Petrunkevitch  1952,  Williams 
1956,  Punzo  2005a)  in  Pepsis  wasps  under 
natural  conditions.  Although  many  aspects 
of  overall  hunting  behavior  for  wasps  of 
this  genus  are  similar  across  species,  the 
results  of  this  field  study  indicate  that  there 
is  some  interspecific  variation  associated 
with  certain  behavioral  components  of 
hunting  between  P.  grossa  and  P.  thisbe 
which  include  frequency  of  drinking  be- 
havior and  sting  insertion  sites.  At  the 
Tornillo  Flat  (TF)  study  site,  females  of 
both  species  approached  a  burrow  occu- 
pied by  adult  females  of  A.  steindachneri 
and  tapped  their  antennae  along  the  edges 
of  the  opening  (burrow  antennation,  BA). 
Wasps  then  entered  the  burrow  after 
cutting  through  the  silk  covering  and 
forced  the  spider  out  onto  the  ground 
surface  (eviction  behavior,  EVB).  These 
behaviors  have  been  reported  for  encoun- 
ters between  theraphosid  spiders  and  other 
species  of  Pepsis  wasps  (Buckley  1862, 
Petrunkevitch  1926,  1952,  Passmore  1936, 
Punzo  2005a),  as  well  as  P.  grossa  (as 
formosa)  (Punzo  and  Garman  1989,  Punzo 
1991)  and  P.  thisbe  (Punzo  1994b),  that  were 
staged  in  the  laboratory. 

Because  A.  steindachneri  is  the  only 
species  of  theraphosid  spider  known  to 
occur  at  TF,  all  females  of  P.  grossa  and  P. 
thisbe  utilized  this  species  as  a  host  for  their 
larvae.  At  sites  25-80  km  to  the  north,  these 
two  species  of  wasps,  along  with  P.  mildei 
Stal,  are  known  to  utilize  both  sexes  of  two 
other  theraphosids,  Aphonopelma  (as  Rhe- 
chostica)  hentzi  and  Dugesiella  (as  Aphono- 
pelma)  echina  Hentzi  as  a  host  (Punzo  and 
Garman  1989,  Punzo  1991).  In  southern 
Texas  (Hidalgo  County),  P.  grossa  (as 
formosa)  utilizes  both  sexes  of  the  thera- 
phosids A.  harlingeninn  (Chamberlin)  and 
A.  heterops  (Chamberlin)  as  hosts,  although 
it  shows  a  strong  preference  for  A.  harlin- 


geninn (Punzo  2006b).  Farther  to  the  west, 
in  Arizona  and  California,  P.  grossa  is 
known  to  hunt  another  theraphosid,  A. 
chalcodes  (Chamberlin)  (Cazier  and  Mor- 
tenson 1964).  In  southern  California,  P. 
thisbe  hunts  two  theraphosid  species,  A. 
reversum  Simon  and  Eurypelma  (as  Aphono- 
pelma) eutylenum  (Ausserer)  (Williams 
1956).  From  the  same  region,  P.  mildei  has 
been  reported  to  utilize  females  of  the 
trapdoor  spider  Bothriocyrtum  californicum 
(Chamberlin  and  Ivie)  (Passmore  1933), 
a  mygalomorph  spider  from  an  entirely 
different  family  (Ctenizidae).  However,  I 
have  never  found  a  nest  containing  any 
species  of  paralyzed  trapdoor  spider  with 
a  Pepsis  larvae,  or  egg  at  any  of  a  number  of 
locations  in  BBNP  or  Big  Bend  Ranch  State 
Park  (Presidio  Co.,  Texas)  (Punzo  unpubl. 
data). 

These  various  host  records  suggest  that 
Pepsis  wasps  have  the  ability  to  utilize 
immatures  and  adult  males  and  females  of 
a  variety  of  theraphosid  spiders  as  hosts, 
depending  on  the  theraphosids  available  at 
any  particular  site.  Presumably,  during  the 
course  of  evolution  in  pompilid  wasps, 
selection  favored  a  preference  for  hunting 
a  single,  larger  host  for  each  wasp  larva. 
Many  species  of  spider  wasps  in  this  genus 
are  among  the  largest  wasps  in  the  New 
World  (Hurd  1952,  Vardy  2000,  2002)  and 
adults  develop  from  larvae  that  attain 
lengths  in  excess  of  27  mm  and  can  weigh 
over  5  g  (Punzo  2000).  In  order  for  a  female 
to  provide  an  adequate  amount  of  food  for 
such  large  larvae,  each  larvae  would  have 
to  be  provided  with  a  high  number  of 
smaller  hosts  either  together  or  over 
a  continuous  period  of  time  (progressive 
provisioning)  as  opposed  to  providing  all 
required  food  at  one  time  (a  single  large 
host  or  mutliple  small  ones,  e.g.  by 
Trypoxylon  wasps)  at  one  time  {mass  pro- 
visioning) (O'Neill  2001).  There  are  obvious 
trade-offs  that  are  involved.  Progressive 
provisioning  involving  several  smaller 
prey  would  require  more  energy  and 
increase  exposure  of  wasps  to  potential 


306  Journal  of  Hymenoptera  Research 

predators,  while  requiring  more  handling  FAPs,  which  are  characteristic  of  instinc- 

time  but  less  risk  from  the  prey.  In  contrast  tive   (innate)   behavior   (Tinbergen    1951). 

a  single,  larger,  more  formidable  host,  may  More  recent  studies,  based  on  sequential 

require  less  handling  time  but  pose  a  higher  laboratory-staged  contests  between  Pepsis 

risk  from  the  prey,  whilst  requiring  less  wasps  and  spiders,  have  shown  that  the 

overall  energy  expenditure  and  a  decreased  amount  of  time  required  to  perform  some 

probability  of  encountering  a  predator.  behavioral  acts  decreases  as  a  function  of 

The  data  collected  to  do  not  point  to  increasing  number  of  encounters  (experi- 

specific  niche  divisions  between  P.  grossa  ence)   (Punzo  and  Garman   1989,  Punzo, 

and  P.  ihisbe  at  the  TF  site.  I  did  not  observe  1991,  2005a). 

any  significant  interspecific  differences  in  The  term  'modal  action  pattern'  (MAP) 

the  sex  or  size  of  spiders  selected,  nor  in  has  been  used  to  describe  components  of 

temporal   patterns    of   hunting   activity,  innate  behavior  that  exhibit  some  degree  of 

Adult   wasps   of  both    species   begin    to  plasticity  (Barlow  1977).  The  acts  that  have 

emerge  from  their  nests  during  late  March  been  shown  to  improve  with  experience 

and    continue    to    do    so    until    August,  include  initial  approach  and  antennation 

Females   of  both   species   were   observed  (AA)  and  attack  and  paralysis  (AP)  (Ta- 

hunting    during    daylight   and    evening  ble  2),  while  the  time  required  for  other 

hours.  At  sites  further  south  (Zapata  Co.,  behavioral   components    does    not.    This 

Texas),  P.  thisbe  begins  to  emerge  in  mid-  suggests  that  some  degree  of  learning  is 

March,  whereas  P.  grossa  adults  are  not  associated  with  hunting  behavior  in  these 

seen  until   mid-April   (F.   Punzo   unpubl.  wasps  (Punzo  1996).  It  is  interesting  to  note 

data).   Perhaps   host  spiders   occurred   at  that  those  acts  which  a  wasp  can  perform 

sufficiently    high    densities    during    the  more  quickly  with  experience  are  precisely 

course  of  the  present  study  at  the  TF  site  those   which   present   the   most   risk   for 

so  that  any  need  for  resource  paritioning  a  wasp.  AA  requires  that  a  wasp  approach 

was  reduced.  within  a  close  distance  of  the  spider  and 

Future  studies  should   further  analyze  actually  touch  the  spider's  body  with  its 

host  preference  in  Pepsis  wasps.  For  exam-  antennae,  placing  it  well  within  the  strike 

pie,  at  locations  where  a  Pepsis  wasp  is  distance  of  the  spider  (Punzo  2007b).  AP 

known  to  utilize  more  than  one  therapho-  requires  the  wasp  to  move  directly  under 

sid    species,   it   would   be   instructive   to  a   spider's   body,    often   passing   directly 

assess  any  differences  that  may  occur  with  below  its  fangs,  grasp  a  leg,  and  insert  its 

respect  to  duration  of  embryonic  develop-  sting.  In  contrast,  performance  of  behav- 

ment,  larval  growth  rate  and  number  of  ioral  acts  that  pose  no  risk  because  the 

larval   instars,   size   of   emerging   adults,  spider    has    already    been    immobilized 

adult  longevity,   fecundity,   flight  endur-  (MG2),  or  are  most  likely  subject  to  bio- 

ance,  and  host  preference,  for  wasps  de-  mechanical  constraints  (BO,  and  BC),  do 

veloping  on  different  hosts.  not  'improve'  with  experience  (Punzo  1991, 

Overall  hunting  behavior  of  Pepsis  wasps  1994b).   Similarly,   MG1,   which   increases 

(P.  grossa,  P.  tliisbe,  P.  mildei,  P.  marginata,  the   distance   between   protagonists   and 

and  Pompilus  spp).  can  be  categorized  into  thereby  decreases  risk  for  a  wasp,  is  not 

several   distinct   behavioral   components  performed  more  rapidly  with  increasing 

(Table  2),  one  of  which  may  or  may  not  number  of  encounters  (Punzo  and  Garman 

occur  (DB)  (Petrunkevitch  1926,  Passmore  1989,  Punzo  2000). 

1936,  Evans  1953).  These  behavioral  com-  During    attack,    these    wasps    showed 

ponents  were  traditionally  interpreted  as  a  marked  preference  for  grasping  legs  3 

examples  of  genetically-determined,  inflex-  or  4  of  a  spider  before  stinging  it.  Perhaps 

ible  acts  referred  to  a  fixed  action  patterns,  the  positions  of  legs  3  and  4,  relative  to  the 


m 


Volume  16,  Number  2,  2007 


307 


center  of  gravity  for  a  spider,  make  it  easier 
for  a  wasp  to  obtain  the  leverage  required 
to  insert  its  sting  through  an  appropriate 
site  on  the  spider's  ventral  surface  in  the 
least  amount  of  time.  The  faster  a  wasp 
insert  its  sting  the  more  rapidly  it  can 
immobilize  a  formidable  host  and  reduce 
the  probability  of  retaliation  by  the  spider. 
Experiments  where  a  wasp  is  presented 
with  a  spider  whose  legs  3  or  4  have  been 
removed,  would  force  a  wasp  to  grasp  leg 
1  or  2,  or  refuse  to  attack  at  all,  and  would 
provide  a  way  to  assess  any  possible 
biomechanical  advantage  associated  with 
grasping  various  legs. 

All  observations  of  encounters  between 
Pepsis  wasps  and  theraphosids  have  shown 
that  a  wasp's  sting  is  directed  into  the 
ventral  body  region  of  a  spider  (Punzo 
2007b,  and  references  cited  therein).  Fe- 
males of  P.  grossa  and  P.  thisbe  showed 
a  marked  preference  when  choosing  a  site 
on  the  spider's  body  in  which  to  insert 
their  sting.  Little  information  is  available 
for  sting  insertion  sites  for  Pepsis  wasps. 
Petrunkevitch  (1926)  observed  a  female  of 
P.  marginata  insert  her  sting  between  the 
third  and  fourth  right  coxae  when  attack- 
ing the  theraphosid,  Cyrtopholis  portoricae 
Simon.  Another  wasp  inserted  its  sting  into 
the  intersegmental  membrane  between  the 
sternum,  maxilla  and  coxa  1.  In  an  encoun- 
ter with  the  theraphosid  Dugesielln  hentzi, 
a  Ptysis  wasp  of  undetermined  species 
inserted  its  sting  through  the  membrane 
between  coxa  3  and  coxa  4  (Baerg  1958). 

The  site  preferred  by  females  of  P.  grossa 
and  P.  thisbe  was  the  intersegmental  mem- 
brane between  coxa  2  and  sternum,  and 
between  coxa  1  and  sternum,  respectively. 
These,  as  well  as  the  other  sting  insertion 
sites  observed  in  this  study  (membrane 
between  pedipalp  and  sternum,  P.  grossa; 
junction  between  abdomen  and  cephalo- 
thorax,  P.  thisbe),  all  allow  a  wasp  to  deliver 
its  venom  into  the  prosomal  nerve  mass 
which  supplies  motoneurons  to  muscles 
involved  in  movments  of  all  legs,  chelicer- 
ae,  and  fangs  (Foelix  1996,  Punzo  2007b). 


Some  investigators  have  observed  in- 
stances in  which  a  wasp  failed  to  locate 
an  insertion  site  during  its  first  attack  on 
a  spider,  moved  a  short  distance  away,  and 
then  attacked  again,  successfully  paralyz- 
ing its  host  (Petrunkevitch  1952,  Williams 
1956,  Baerg  1958).  Nonetheless,  once  the 
sting  delivers  venom  into  the  prosomal 
nerve  mass,  paralysis  of  the  spider  occurs 
very  rapidly  as  indicated  by  a  curling  of 
the  spider's  legs  under  its  body,  slight 
twitching  movements  of  some  appendages, 
and  then  complete  immobilization  (Petrun- 
kevitch 1926,  Cazier  and  Mortenson  1964, 
Punzo  2000). 

These  results  on  sting  insertion  sites  also 
indicate  that  there  is  behavioral  variation 
exhibited  by  these  wasps.  This  suggests 
several  interesting  questions  that  future 
studies  should  address:  do  individual 
wasps  choose  the  same  insertion  site  for 
all  encounters,  or  do  they  vary?  If  sting 
insertion  sites  are  'fixed'  for  individual 
females  this  would  suggest  that  the  behav- 
ioral program  has  a  genetic  basis  (innate). 
If  so,  breeding  experiments  involving 
males  with  females  showing  different 
behavioral  phenotypes  might  shed  some 
light  on  the  patterns  of  inheritance  in- 
volved in  this  behavior.  Secondly,  is  there 
a  relationship  between  a  particular  inser- 
tion site  and  time  required  to  immobilize 
a  spider? 

Nutritional  state  (body  condition)  may 
afford  a  possible  explanation  for  why 
drinking  behavior  (DB)  occurs  in  only 
some  encounters.  Drinking  hemolymph 
oozing  from  a  host's  wound  site  may 
provide  necessary  nutrients  to  meet  the 
energetic  demands  of  flight  which  wasps 
engage  in  when  searching  for  hosts,  as  well 
as  those  of  venom  production,  handling 
time,  and  burrow  closure.  Drinking  fluids 
from  a  spider's  mouth  cavity  may  help 
wasps  to  maintain  proper  water  balance  of 
body  fluids.  Similar  behavior  has  been 
reported  for  species  in  other  wasp  families. 
For  example,  Tinbergen  (1972)  observed 
that  females  of  the  digger  wasp,  Philanthus 


308 


Journal  of  Hymenoptera  Research 


triangulum  Fabricius,  which  selectively 
hunt  honeybees,  Apis  mellifera  L.,  press 
the  abdomen  of  a  paralyzed  bee  through 
their  mandibles  and  lick  up  the  fluid 
(nectar)  extruded  from  the  bee's  mouth. 
Species  of  spider  wasps  from  other 
genera  are  also  known  to  drink  fluids  from 
a  host's  mouth  cavity  or  wound  site 
(Petrunkevitch  1952,  Evans  1953,  Williams 
1956,  Evans  and  West-Eberhard  1970, 
Punzo  2000). 

It  may  be  that  wasps  engage  in  DB  only 
after  a  certain  number  of  foraging  bouts 
have  occurred  resulting  in  a  need  to 
replenish  nutrients  and /or  body  fluids. 
Experimental  protocols  using  a  tethered 
flight  apparatus  should  be  used  in  future 
studies  to  test  this  hypothesis.  Different 
female  wasps  could  be  subjected  to  forced 
flight  tests  for  varying  periods  of  time  and 
then  allowed  to  encounter  a  host.  If  the 
hypothesis  is  true,  wasps  that  are  subjected 
to  longer  bouts  of  flight  (and  thus  expend 
more  energy,  and  lose  more  water  by 
evaporation)  should  be  more  likely  to 
engage  in  DB  than  wasps  subjected  to 
flying  for  shorter  periods  of  time.  Such  an 
experimental  design  would  also  allow  one 
to  determine  the  amount  of  flight  time 
required  to  initiate  LB  in  a  particular  wasp 
species. 

The  confines  of  a  spider's  burrow  might 
not  provide  enough  room  for  a  wasp  to 
maneuver  in  such  a  way  as  to  effectively 
administer  a  sting  to  its  host.  This  may 
account  for  the  fact  that  all  wasps  observed 
at  the  TF  site  forced  a  spider  out  of  its 
burrow  and  onto  the  ground  surface 
(eviction  behavior,  EVB)  before  attacking 
their  host.  Similar  EVB  has  been  observed 
for  Pepsis  wasps  presented  with  tarantulas 
that  have  been  allowed  to  excavate  bur- 
rows within  their  cages  under  laboratory 
conditions  (Petrunkevitch  1926,  Punzo  and 
Garman  1989,  Punzo  1991,  1994b). 

Interspecific  differences  in  the  amount  of 
time  required  to  complete  the  overall 
hunting  sequence  among  P.grossa  and  P. 
thisbe  in  the  field  may  be  associated  with 


some  wasps  of  either  species  having  had 
more  encounters  with  hosts  than  other 
wasps.  It  may  also  reflect  genetically-based 
differences  in  synaptic  events  associated 
with  afferent  (sensory)  neural  pathways 
involved  with  detection  and  identification 
of  hosts  and /or  efferent  (motor  pathways) 
involved  in  the  control  of  bodily  move- 
ments required  for  various  behavioral  acts. 

On  a  final  note,  theraphosid  spiders  are 
typically  aggressive  and  innately  strike  at 
arthropods  that  wander  within  their  prey 
awareness  area  (Punzo  2007b).  The  fact 
that  no  spider  attempted  to  seize  a  wasp 
suggests  that  the  spider's  attack  response  is 
somehow  inhibited.  It  has  been  suggested 
that  these  wasps  may  release  some  chemi- 
cal compound (s)  or  possess  chemosensory 
cues  associated  with  their  epicuticle  that 
inhibit  spider's  from  attacking  them  (Pet- 
runkevitch 1952,  Punzo  2000).  Petrunke- 
vitch (1926)  observed  that  P.  marginata 
from  Puerto  Rico  produced  a  "pungent 
odor"  when  initially  making  contact  with 
a  theraphosid  host  and  argued  that  the 
substance  responsible  for  this  odor  might 
somehow  diminish  the  aggressiveness  of 
the  spider.  Others  have  pointed  out  that 
the  smooth  surface  of  a  wasp's  cuticle, 
combined  with  its  high  degree  of  hardness, 
makes  it  difficult  for  a  spider's  fangs  to 
penetrate  a  wasp's  integument  (Petrunke- 
vitch 1926,  Passmore  1936). 

It  should  be  pointed  out  that  Petrunke- 
vitch (1926)  observed  a  theraphosid  (Cyrto- 
pholis  portoricae  Simon)  that  unsuccessfully 
attempted  to  grasp  a  female  of  P.  marginata 
Lucas  with  her  fangs  as  the  wasp  passed 
under  the  spider.  Cazier  and  Mortenson 
(1964)  observed  a  Pepsis  grossa  (as  formosa) 
female  entering  a  burrow  occupied  by 
Aphonopelma  sp.,  and  after  a  few  minutes 
the  spider  emerged  from  its  burrow  with 
its  anterior  two  legs  wrapped  around  the 
wasp  and  its  chelicerae  inserted  into  the 
wasp's  abdomen.  Nonetheless,  the  wasp 
was  able  to  sting  the  spider.  After  being 
stung,  the  spider  released  the  wasp  which 
exhibited  erratic  movements  and  was  un- 


Volume  16,  Number  2,  2007 


309 


able  to  fly.  The  spider's  right  leg  was 
rigidly  extended  forward,  making  locomo- 
tion awkward.  After  several  minutes,  the 
wasp  and  spider  were  placed  in  a  screened 
plastic  container  and  initially  both  animals 
avoided  one  another.  When  observed 
45  min  later,  the  spider  was  engaged  in 
eating  the  wasp.  These  observations  sug- 
gest that:  (1)  cues  potentially  responsible 
for  inhibiting  a  spider's  strike  response 
may  not  always  be  effective  (2)  varying 
degrees  of  effectiveness  may  be  species- 
specific;  or  (3)  mutations  may  account  for 
differences  in  the  chemical  profile  of  the 
wasp's  cuticle  and  certain  profiles  may  be 
less  effective  at  deterring  a  spider's  strike 
than  others. 

ACKNOWLEDGMENTS 

I  thank  C.  Bradford,  L.  Ludwig,  G.  Broad,  and 
anonymous  reviewers  for  commenting  on  an  earlier 
draft  of  the  manuscript,  and  L.  Ludwig,  J.  Bottrell,  K. 
Crawford,  and  S.  Madragon  for  assistance  in  field 
observations.  B.  Garman  generously  provided  consul- 
tation on  statistical  analyses.  This  research  was 
supported  by  a  Faculty  Development  Grant  from  the 
University  of  Tampa.  Field  work  was  conducted  with 
permission  of  the  National  Park  Service,  with  logistical 
support  provided  by  R.  Skiles,  Big  Bend  National  Park. 

LITERATURE  CITED 

Baerg,  W.  J.  1958.  The  Tarantula.  University  of  Kansas 
Press,  Lawrence,  Kansas.  88  pp. 

Barlow,  G.  W.  1977.  Modal  action  patterns.  Pp.  98-134 
in:  T.  A.  Seboek,  ed.  How  Animals  Communicate. 
Indiana  Univ  Press,  Bloomington,  Indiana. 

Buckley,  S.  B.  1862.  The  tarantula  (Mygale  hentzii 
Girard)  and  its  destroyer  {Pompilus  formosus  Say). 
Proceedings  of  the  Entomological  Society  of  Philadel- 
phia 1:  138-139. 

Cazier,  M.  A.  and  M.  Mortenson.  1964.  Bionomical 
observations  on  tarantula  hawks  and  their  prey 
(Hymenoptera:  Pompilidae):  Pepsis.  Annals  of  the 
Entomological  Society  of  America  57:  533-541. 

Evans,  H.  E.  1953.  Comparative  ethology  of  spider 
wasps.  Systematic  Zoology  2:  155-172. 

and  M.  J.  West-Eberhard.   1970.   The  Wasps. 

University  of  Michigan  Press,  Ann  Arbor,  Michi- 
gan, vii  +  265  pp. 

Foelix,  R.  F.  1996.  The  Biology  of  Spiders.  Oxford 
University  Press,  London,  v  +  321  pp. 

Gabel,  J.  1972.  Further  observations  of  theraphosid 
tarantula  burrows.  Pan-Pacific  Entomologist  48: 
72-73. 


Haupt,  H.  1952.  Alte  und  neue  Pepsis  Arten  (Hyme- 
noptera-Sphecoidea).  Nova  Acta  Eeopoldina  N.F. 
15:  309-414. 

Hurd,  P.  D.  1952.  Revision  of  the  Nearctic  species  of 
the  pompilid  genus  Pepsis  (Hymenoptera:  Pom- 
pilidae). Bulletin  of  the  American  Museum  of  Natural 
History  98:  257-334. 

Ito,  Y.  1980.  Comparative  Ethology.  Second  Ed.  Cam- 
bridge University  Press,  Cambridge.  478  pp. 

Kurczewski,  F.  E.  and  E.  J.  Kurczewski.  1968.  Host 
records  for  some  North  American  Pompilidae 
(Hymenoptera)  with  a  discussion  of  factors  in 
prey  selection.  Journal  of  the  Kansas  Entomological 
Society  41:  1-33. 

Minch,  E.  1979.  Annual  activity  patterns  in  the 
tarantula  Aphonopelma  chalcodes  Chamberlin.  No- 
vitates  Arthropodae  1:  1-34. 

Nation,  J.  L.  2002.  Insect  Physiology  and  Biochemistry. 
CRC  Press,  Boca  Raton,  Florida.  485  pp. 

Parent,  L.  1996.  Big  Bend  National  Park.  Falcon 
Publishing,  Helena,  Montana,  viii  +  171  pp. 

Passmore,  L.  1933.  California  trapdoor  spider  per- 
forms engineering  marvels.  National  Geographic 
Magazine  64:  195-211. 

.  1936.  Tarantula  and  tarantula  hawk.  Nature 

Magazine  27:  155-159. 

Petrunkevitch,  A.  1926.  Tarantula  vs.  tarantula  hawk: 
a  study  in  instinct,  journal  of  Experimental  Zoology 
43:  367-397. 

.    1952.   The   spider   and    the   wasp.   Scientific 

American  187:  20-33. 

Powell,  A.  M.  1988.  Trees  and  Shrubs  of  Trans  Pecos 
Texas.  Big  Bend  Natural  History  Association,  Big 
Bend,  Texas.  536  pp. 

Price,  P.  W.  1975.  Strategies  for  egg  production. 
Evolution  28:  76-84. 

.  1997.  Insect  Ecology.  Third  Ed.  J.  Wiley,  New 

York,  vi  +  874  pp. 

Punzo,  F.  1991.  Neurochemical  events  associated  with 
learning  and  hunting  behavior  in  the  spider 
wasp,  Pepsis  formosa  (Hymenoptera:  Pompilidae). 
Florida  Scientist  54:  51-61. 

.  1994a.  The  biology  of  the  spider  wasp,  Pepsis 

thishe  (Hymenoptera:  Pompilidae)  from  Trans 
Pecos  Texas.  I.  Adult  morphometries,  larval 
development  and  the  ontogeny  of  feeding  behav- 
ior. Psyche  101:  229-241. 

.  1994b.  The  biology  of  the  spider  wasp,  Pepsis 

thishe  (Hymenoptera:  Pompilidae)  from  Trans 
Pecos  Texas.  II.  Temporal  patterns  of  activity 
and  hunting  behavior  with  special  reference  to 
effects  of  experience.  Psyche  101:  243-256. 

.    1996.    Localization    of   brain    function    and 

neurochemical  events  associated  with  learning 
in  insects.  Recent  Trends  in  Comparative  Biochemis- 
try and  Physiology  2:  9-16. 

.  2000.  Desert  Arthropods:  I  ife  History  Variations. 


Springer,  Heidelberg,  Germany,  xi  +  310  pp. 


310 


Journal  of  Hymenoptera  Research 


— .  2005a.  Experience  affects  hunting  behavior  of 
the  wasp,  Pepsis  mildei  Stal  (Hymenoptera:  Pom- 
pilidae).  journal  of  the  New  York  Entomological 
Society  113:  222-229. 

— .  2005b.  The  ability  of  Aphonopelma  steindachneri 
(Ausserer)  (Arachnida:  Theraphosidae)  to  detect 
and  respond  to  chemosensory  cues  associated 
with  a  predator,  Hogna  carolinensis  (Arachnida: 
Lycosidae).  Bulletin  of  the  British  Arachnological 
Society  13:  169-172. 

— .  2005c.  Studies  on  the  natural  history,  ecology, 
and  behavior  of  Pepsis  cerberus  and  P.  mexicana 
(Hymenoptera:  Pompilidae)  from  Big  Bend  Na- 
tional Park,  Texas,  journal  of  the  New  York 
Entomological  Society  113:  84-95. 

— .  2006a.  Plants  whose  flowers  are  utilized  by 
adults  of  Pepsis  grossa  Fabricius  (Hymenoptera: 
Pompilidae)  as  a  source  of  nectar.  Journal  of 
Hvmenoptera  Research  15:  171-176. 

— .  2006b.  Effect  of  host  odor  cues  on  behavioral 
responses  of  the  spider  wasp,  Pepsis  formosa 
(Hymenoptera:  Pompilidae).  Texas  journal  of 
Science  58:  45-55. 

— .  2007a.  Microhabitat  utilization,  diet  composi- 
tion, and  diel  periodicity  in  five  sympatric  species 
of  desert  arachnids:  a  wolf  spider  (Hogna  caroli- 
nensis), tarantula  spider  (Aphonopelma  steindach- 
neri), solifuge  (Eremobates  palpisetulosus),  giant 
whipscorpion  (Mastigoproctus  giganteus),  and 
scorpion  (Diplocentrus  bigbendensis).  Bulletin  of 
the  British  Arachnological  Society  14:  1-8. 

— .  2007b.  Spiders:  Biology,  Ecology,  Natural  History 
and  Behavior.  Brill  Academic  Publishers,  Leiden, 
The  Netherlands,  iii  +  431  pp. 

—  and  B.  Garman.  1989.  Effects  of  encounter 
experience  on  the  hunting  behavior  of  the  spider 
wasp,  Pepsis  forniosa  (Say)  (Hymenoptera:  Pom- 
pilidae). South-western  Naturalist  34:  513-518. 

—  and  L.  Henderson.  1999.  Aspects  of  the  natural 
history  and  behavioral  ecology  of  the  tarantula 
spider  Aphonopelma  hentzi  (Chamberlin)  (Orthog- 
natha,  Theraphosidae).  Bulletin  of  the  British 
Arachnological  Society  11:  121-128. 


and  L.  Ludwig.  2005.  Behavioral  responses  of 

Pepsis  thisbe  (Hymenoptera:  Pompilidae)  to  che- 
mosensory cues  associated  with  host  spiders. 
Journal  of  Insect  Behavior  18:  757-766. 

Rau,  P.  and  N.  Rau.  1918.  Wasp  Studies  Afield. 
Princeton  University  Press,  Princeton,  New  Jer- 
sey. 372  pp. 

Schmidt,  J.  O.  2004.  Venom  and  the  good  life  in 
tarantula  hawks  (Hymenoptera:  Pompilidae): 
how  to  eat,  not  be  eaten,  and  live  long,  journal 
of  the  Kansas  Entomological  Society  77:  402-413. 

Smith,  A.  1994.  Tarantula  Spiders:  Tarantulas  of  the 
U.S.A.  and  Mexico.  Fitzgerald  Press,  London. 
312  Pp. 

Sokal,  R.  R.  and  F.  J.  Rohlf.  1995.  Biometry.  Third  Ed. 
W.  H.  Freeman,  New  York.  881  pp. 

Tinbergen,   N.    1951.    The   Study  of  Instinct.   Oxford 
University  Press,  Oxford.  228  pp. 
— .  1972.  The  Animal  in  its  World:  Field  Studies. 
Harvard  University  Press,  Cambridge,  Massachu- 
setts. 343  pp. 

U.  S.  Department  of  Interior.  2005,  Division  of 
Publications,  National  Park  Service,  National 
Park  Handbook,  No.  122,  Washington,  D.C.,  216 

PP- 

Vardy,  C.  R.  2000.  The  New  World  tarantula-hawk 
wasp  genus  Pepsis  Fabricius  (Hymenoptera: 
Pompilidae).  Part  1.  Introduction  and  the  P.  rubra 
species-group.  Zoologische  Verhandelingen  332: 
1-86. 

.  2002.  The  New  World  tarantula-hawk  wasp 

genus  Pepsis  Fabricius  (Hymenoptera:  Pompili- 
dae). Part  2.  The  P.  grossa  to  P.  deaurata  groups. 
Zoologische  Verhandelingen  338:  1-135. 

Vinson,  S.  B.  1984.  Parasitoid-host  relationship.  Pp.  in: 
W.  J.  Bell,  and  R.  T.  Carde  eds.  Chemical  Ecology  of 
Insects.  Sinauer  Associates,  Sunderland,  Massa- 
chusetts 

Williams,  F.  X.  1956.  Life  history  studies  of  Pepsis  and 
Hemipepsis  wasps  in  California  (Hymenoptera: 
Pompilidae).  Annals  of  the  Entomological  Society  of 
America  49:  447-466. 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  311-325 

Systematic  Studies  on  the  Pompilidae  Occurring  in  Japan:  Genus 
Irenangelus  Schulz  (Hymenoptera:  Pompilidae:  Ceropalinae) 

Akira  Shimizu  and  Raymond  Wahis 

(AS)  Department  of  Biological  Sciences,  Graduate  School  of  Science  and  Engineering, 

Tokyo  Metropolitan  University,  Minami-Ohsawa  1-1,  Hachioji, 

Tokyo,  192-0397  Japan;  email:  shimizu-akira@cmetro-u.ac.jp 

(RW)  Entomologie  fonctionnelle  et  evolutive,  Faculte  universitaire  des  Sciences  agronomiques, 

B.  5030  Gembloux,  Belgique;  email:  entomologie@fsagx.ac.be  and  raymond.wahis@skynet.be 


Abstract. — The  Japanese  species  of  the  genus  Irenangelus  Schulz  (Pompilidae:  Ceropalinae)  are 
revised.  Three  new  species  are  described:  I.  hikosamis  Wahis,  /.  nantbui  Shimizu,  and  I. 
punctipleuris  Wahis.  Irenangelus  hikosamis  occurs  in  Japan  (Honshu,  Shikoku,  Kyushu),  Korea, 
Taiwan,  and  China;  J.  nambui  occurs  in  Japan  (Honshu)  and  Korea;  J.  punctipleuris  is  broadly 
distributed  from  Japan  through  the  Philippines  and  Malaysia  to  India  and  Sri  Lanka. 


Irenangelus  Schulz  1906,  like  Ceropales 
Latreille  1796,  is  one  of  the  most  aberrant 
genera  of  the  family  Pompilidae.  All 
members  of  both  genera  have  an  uncoiled 
antenna,  reniform  eyes  with  the  inner 
orbits  strongly  emarginate  and  diverging 
above,  a  fully  exserted  labium  in  both 
sexes,  and  a  strongly  compressed  metaso- 
mal  sternum  VI  produced  beyond  tergum 
VI  in  the  female. 

Evans  (1969,  1987)  reviewed  the  Neo- 
tropical species  of  Irenangelus.  In  these 
papers  he  treated  ten  species,  seven  of 
which  were  new,  and  regarded  Xantham- 
pulex  Schulz  1906  as  a  synonym  of  Irenan- 
gelus. Kimsey  and  Wasbauer  (2004)  revised 
the  New  World  species  of  Irenangelus, 
adding  two  new  species.  As  regards  the 
Old  World  species  of  Irenangelus,  Cameron 
(1891, 1896),  Bingham  (1896),  Schulz  (1906), 
Turner  (1910),  Rohwer  (1919),  Banks 
(1934),  and  Wahis  (1988)  described  either 
single  species  or,  at  most,  a  few  new 
species,  but  no  comprehensive  revisionary 
studies  have  been  published. 

Species  of  Irenangelus  have  long  been 
recognized  as  cleptoparasites  of  other 
pompilids,  as  are  species  of  Ceropales.  Thus, 


Williams  (1919)  reared  an  Irenangelus  wasp 
from  a  nest  of  Auplopus  nyemitawa  (Rohwer 
1919)  and  several  wasps  in  this  genus  from 
cocoons  of  nests  of  Tachypompilus  analis 
(Fabricius  1781)  in  the  Philippines.  At  least 
one  wasp  reared  belonged  to  I.  luzonensis 
(Rohwer  1919).  In  Costa  Rica,  Wcislo  et  al. 
(1988)  observed  females  of  /.  eberhardi 
Evans  1987  fly  or  perch  near  nests  of 
Auplopus  seinialatus  Dreisbach  1963,  enter 
an  open  cell  containing  a  spider,  and 
extend  her  gaster  deep  into  the  cell.  They 
reared  several  wasps  of  this  parasite  from 
one  of  the  nests  of  A.  semialatus.  Shimizu 
(see  below)  found  females  of  a  Japanese 
species  of  Irenangelus  attempting  to  ovipos- 
it eggs  into  a  slit  of  the  booklung  of 
heteropodid  spiders  that  the  host  pompi- 
lids had  captured  and  then  transported. 

Irenangelus  is  mainly  distributed  in  the 
Oriental  and  Neotropical  regions.  In  Japan 
only  one  species  of  the  genus  has  been 
known  since  Yasumatsu  (1933)  recorded 
the  species  from  Honshu,  Kyushu,  and 
Taiwan  as  Xanthampulex  pernix  (Bingham 
1896).  This  species  is  distributed  also  in 
Korea  and  China,  and  differs  from  X. 
pernix,  which  was  originally  recorded  from 


312  Journal  of  Hymenoptera  Research 

"Tenasserim"    (Burma).    Recently   speci-  vis,  California,  USA;  ELKU,  Collection  of 

mens  of  a  further  two  species  of  this  genus  Entomological   Laboratory,   Kyushu   Uni- 

were  collected  from  Honshu,  Japan.  One  of  versify,  Fukuoka,  Japan;  AEIC,  American 

them   is   found   in   Korea   in   addition   to  Entomological  Institute,  Gainesville,  Flor- 

Japan,  and  the  other  occurs  from  Japan  ida,   USA;   FSAG,    Entomologie   fonction- 

through  Southeast  Asia  to  South  Asia.  It  nelle   et   evolutive,   Faculte    universitaire 

was   found   that   these   three   species   are  des   Sciences   agronomiques,   Gembloux, 

undescribed.  Belgique;  RMNH,  Nationaal  Naturhistor- 

In   this  paper,   we  review   the  generic  ische    Museum,     Leiden,     Netherlands; 

characters  and  phylogenetic  relationships  BMNH,  Natural  History  Museum,  London, 

of  Irenangelus,  describe  three  new  species  UK;  OMNH,  Osaka  Museum  of  Natural 

and  provide  a  key  to  their  identification.  History,   Osaka,   Japan;   CNC,   Canadian 

National   Collection   of   Insects,   Ottawa, 

MATERIALS  AND  METHODS  Ontario,  Canada;  NSMT,  Department  of 

The  terminology  of  the  wing  veins  and  Zoology,   National  Science  Museum,  To- 

cells   follows   Day   (1988).   The   following  kyo>  JaPan;  TMUB,  Laboratory  of  Zoolog- 

morphological  terms  and  abbreviations  are  ical  Systematica  Department  of  Biological 

used:  antennocular  line,  the  anterior  mar-  Sciences,  Tokyo  Metropolitan  University, 

gin   of  the   frons   in   dorsal   view;   scutal  Tokyo,  Japan;  USNM,  National  Museum  of 

groove,   a   pair   of   longitudinal    grooves  Natural  History,  Washington,  D.  C,  USA. 

between  the  notaulus  and  parapsidal  sul- 

cus  on  the  mesoscutum  (Evans  (1969)  and 

Kimsey  and  Wasbauer  (2004)  called  this  ^           F               ,      c  ,     , 

y      ,     „.   _  T_    t    \           .              ,  Genus  Irenangelus  Schulz 
the    notaulus  );  LID,  the  lower  interocular 

distance;    MID,    the    middle    interocular  Irenangelus  Schulz  1906:   175.  Type  of  genus: 

distance;    OOL,    the    ocello-ocular    line;  irenangelus  hornus  Schulz  1906: 160,  by  mono- 

POL,  the  postocellar  line;  SMC,  the  sub-  v    \         ,     c  ,    ,    1fin^  1DO  T          , 

.    r ..      .    .       .            .         TTTT^     ,  Xanthampulex  Schulz  1906:  1 83.  Type  of  genus: 

marginal  cell  of  the  tore  wing;  UID,  the  v     ,,          ,       ,   c      c  ,    ,      /ani-     1QQ    u 

&                                                   &/           '  Xmitluimpulex    tnfur   Schulz    1906:    183,    by 

upper  interocular  distance.  monotvpv 

Measurements  were  made  in  the  follow- 
ing ways:  clypeus  length  versus  breadth,  Description. — Further  to  the  descriptions 
being  measured  comparing  the  length  of  of  this  genus  by  Evans  (1969)  and  Kimsey 
the  clypeus  from  the  uppermost  point  of  and  Wasbauer  (2004)  we  note  the  following 
the  front-clypeal  sulcus  to  the  apical  characteristics:  gena  flattened  or  concave 
margin  to  the  maximum  breadth  of  the  just  posterior  to  outer  orbit  at  least  below; 
clypeus;  labrum  length  versus  breadth,  posterolateral  margin  of  pronotum  almost 
being  measured  comparing  the  longest  straight  (Figs  2F,  3D);  scutal  groove  deeply 
part  of  the  labrum  to  the  breadth  across  impressed  in  many  species  (Figs  1A,  2C); 
the  base  of  the  labrum;  breadth  of  flagello-  metapostnotum  well  developed  at  least 
mere  I,  being  measured  across  the  maxi-  medially  (Figs  1A,  D,  2 A,  F,  3D);  fore  wing 
mum  breadth  of  flagellomere  I  in  dorsal  vein  M  reaching  outer  wing  margin 
view.  In  the  description  of  each  species,  the  (Figs  2A,  5A,  B);  metatibia  with  longitudi- 
measurements  of  the  holotype  are  given  in  nal  sharp  groove  along  upper  margin  of 
parentheses.  brush  on  inner  side  (Fig.  2A);  apicoventral 

Specimen  depositories  are  abbreviated  seta  on  metatarsomere  V  long  and  setiform 

as   follows:   ZMUC,   Zoological   Museum,  (see  Shimizu   et  al.    1998:   fig.   3);   female 

University  of  Copenhagen,  Copenhagen,  laterosterna    of   metasomal    sternum    VI 

Denmark;  UCDC,  R.  M.  Bohart  Museum  of  extending  dorsad,  scarcely  overlapping  to 

Entomology,  University  of  California,  Da-  envelope   sting   apparatus;   sting   almost 


Volume  16,  Number  2,  2007 


313 


Fig.  1.  Irenangelus  hikosanus  n.  sp.  (A-F,  holotype  female;  G-I,  para  type  male  from  Japan)  and  female  /.  pernix 
Bingham  from  Bali,  Indonesia  (J).  A,  Head  and  mesosoma,  dorsal  view;  B,  head,  lateral  view;  C,  head,  frontal 
view;  D,  mesosoma,  lateral  view;  E,  mesosternum  and  mesocoxa,  ventrolateral  view;  F,  right  metatarsal  claw, 
outer  view;  G,  genitalia  (left  half,  ventral  view;  right  half,  dorsal  view);  H,  subgenital  plate,  ventral  view;  I, 
sternum  VI,  ventral  view.  Scale  lines:  0.5  mm. 


straight;  male  sternum  VI  without  a  pair  of 
sublateral  hook-like  projections  posteriorly 
(Fig.  II);  digitus  volsellaris  with  large 
semicircular  emargination  on  inner  margin 
(Figs  1G,  21,  3L).  ' 

Diagnosis. — Irenangelus  is  closely  related 
to  Ceropales,  forming  a  monophyletic 
group,  the  subfamily  Ceropalinae  (Shimizu 
1994,  Pitts  et  al.  2006).  These  two  genera  are 
distinguishable  on  the  characteristics 
shown  in  Table  1. 

Phylogenetic  relationships. — On  the  basis 
of  morphological  characteristics,  both  Shi- 
mizu (1994)  and  Pitts  et  al.  (2006)  treated 


the  Ceropalinae  (Ceropales  +  Irenangelus)  as 
the  most  basal  clade  in  the  Pompilidae.  This 
subfamily  is  thus  considered  to  be  the  sister 
group  to  the  rest  of  the  family.  Based  on  this 
hypothesis,  there  is  a  biological  inconsisten- 
cy: although  the  Ceropalinae  are  placed  as 
the  earliest  offshoot  of  the  pompilid  stock, 
all  species  of  the  subfamily  whose  behav- 
iour is  known  are  cleptoparasitic.  To  avoid 
this  contradiction,  Shimizu  (1994)  consid- 
ered that  the  behavioural  type  of  the 
Ceropalinae  has  evolved  directly  from  that 
of  the  parasitoids,  which  is  the  most  likely 
life  history  of  a  common  ancestor  of  the 


314 


Journal  of  Hymenoptera  Research 


Table  1.     Comparison  of  differential  characters  between  Irenangelus  and   Ceropales  in  the  broad   sense, 
including  Priesnerius  and  Hemiceropales. 


Irenangelus 


Ceropales 


1.     Metacoxa  normal-sized,  less  than  1.5X  as  long  as    Metacoxa  much  larger  than  mesocoxa,  more  than  1.5X 


mesocoxa. 


as  long  as  mesocoxa. 


2.     Ventral  angle  of  pronotum  short  and  blunt,  not        Ventral  angle  of  pronotum  long  and  acute,  partly 


attaining  dorsal  margin  of  procoxa  (Fig.  3D). 

3.  Posterolateral  margin  of  pronotum  almost 
straight  (Figs  2F,  3D). 

4.  Inner  margin  of  male  protarsomere  V  not 
produced. 

5*.     Male  subgenital  plate  flat  or  gently  convex 
(Figs  1H,  2H,  3L). 


covering  dorsal  margin  of  procoxa. 
Posterolateral  margin  of  pronotum  strongly  curved 

inward. 
Inner  margin  of  male  protarsomere  V  produced 

ventrally. 
Male  subgenital  plate  tectate  with  median  carina. 


*  Character  treated  by  Kimsey  and  Wasbauer  (2004). 

Pompilidae.  Similarly,  Day  (1988: 16)  stated: 
"it  seems  more  probable  that  Ceropales  has 
evolved  from  an  ancestral  group  already 
specialized  as  ectoparasitioids."  This  pre- 
sumption seems  likely  given  that  the  two 
important  characteristics  of  Ceropalinae 
females  are  shared  with  other  pompilids 
known  to  behave  as  parasitoids,  viz-  (i)  the 
strongly  exposed  clypeus,  and  (ii)  the 
laterally  compressed  sternum  VI. 

Biology. — One  or  more  Philippine  species 
of  Irenangelus  are  known  to  be  cleptopar- 
asites  of  Auplopus  nyemitawa  (Pepsinae) 
and  Tachypompilus  analis  (Pompilinae), 
and  I.  eberlmrdi  a  cleptoparasite  of  Auplopus 
semialatus  (Pepsinae)  (Williams  1919).  Shi- 


mizu  observed  the  cleptoparasitic  behav- 
iour of  /.  lukosanus  Wahis  n.  sp.,  which 
pursues  its  host  pompilid  Platydialepis 
ryoheii  (Ishikawa  1956)  (Pepsinae)  as  the 
latter  transports  her  prey,  Heteropoda  for- 
cipata  (Karsch  1881)  (Heteropodidae)  to  her 
nest.  Eventually  the  parasite  pounces  on 
the  spider  and  extends  her  gaster,  attempt- 
ing to  insert  its  tip  into  a  slit  of  the  spider's 
booklung.  The  details  of  this  behaviour 
will  be  treated  in  a  separate  paper. 

Distribution. — This  genus  is  known  from 
Oriental,  Neotropical,  Australian,  East 
Asian,  and  Madagascan  Regions  (see  Wa- 
his 1988),  but  is  best  represented  in  the  first 
two  regions. 


KEY  TO  FEMALES  AND  MALES  OF  IRENANGELUS  OCCURRING  IN  JAPAN 

Flagellum  crenulate  in  profile,  i.e.,  flagellomeres  II— X  each  with  angular  swelling 
below  (Fig.  3A);  all  tarsal  claws  bifid,  inner  ray  truncate  (Fig.  3G);  propodeum 
transversely  striate;  metasomal  tergum  I  gradually  narrowed  and  petiolate  (tergum 
I  narrower  immediately  behind  articulation  with  propodeum  than  width  at 
articulation  itself)  (Fig.  3E)  or  parallel-sided  basally;  head  and  mesosoma  with 
irregularly-distributed  punctures  (Figs  4C-F).  (Head  and  mesosoma  black,  varie- 
gated with  bright  yellow  markings;  metasoma  and  legs  predominantly  reddish 
brown;  body  length  8-12  mm) I.  punctipleuris  Wahis,  n.  sp. 

Flagellum  not  crenulate  in  profile;  all  tarsal  claws  dentate  (Fig.  IF)  or  sub-bifid 
(Fig.  2G),  inner  ray  acute;  propodeum  smooth,  never  striate;  metasomal  tergum  I 
abruptly  narrowed,  not  petiolate  or  parallel-sided  basally  (Fig.  2A);  head  and 
mesosoma  impunctate  (Fig.  4B) 2 

Propodeum  with  lateral  tubercle  between  spiracle  and  posterior  rim  (Fig.  1A); 
interantennal  area  distinctly  raised  (Figs  IB,  4B);  fore  wing  crossvein  cu- 
a  originating  at  or  slightly  distal  to  separation  of  vein  M+Cu  (Fig.  5A);  fore  wing 


Volume  16,  Number  2,  2007 


315 


SMC3  removed  by  approximately  its  own  length  from  outer  wing  margin;  apical 
margin  of  subgenital  plate  strongly  convex  (Fig.  1H);  body  predominantly 
yellowish  brown;  body  length  8-15  mm    J.  hikosanus  Wahis,  n.  sp. 

Propodeum  without  lateral  tubercle  between  spiracle  and  posterior  rim  (Fig.  2A); 
interantennal  area  not  raised,  continuous  to  upper  frons  (Fig.  2D);  fore  wing 
crossvein  cu-a  originating  slightly  basad  of  separation  of  vein  M+Cu  (Fig.  2A);  fore 
wing  SMC3  removed  by  much  more  than  its  own  length  from  outer  wing  margin; 
apical  margin  of  subgenital  plate  slightly  emarginate  or  almost  truncate  (Fig.  2H); 

body  predominantly  blackish  brown  to  black;  body  length  5-9  mm    

I.  ttambui  Shimizu,  n.  sp. 


Irenangelus  hikosanus  Wahis,  new  species 

(Figs  1A-I,  4B,  5A) 

Xanthampulex  pernix:  Yasumatsu  1933:  143, 
figure  1,  Jo,  misidentification;  Kim  1970:  807. 

Irenangelus  pernix:  Lelej  et  al.  1994:  145;  Lelej  et 
al.  1995:  46;  Shimizu  1994:  45;  Shimizu  1996: 
507;  Shimizu  et  al.  1998:  429,  figure  3. 

Female. — Length:  Body  9.1-14.5  (9.8)  mm; 
fore  wing  8.7-11.3  (8.7)  mm.  Coloration: 
Body  and  appendages  predominantly  yel- 
lowish brown  and  polished.  Following 
light  yellow:  mandible  (apical  portion  dark 
brown),  clypeus  (lateral  side  sometimes 
yellowish  brown),  frons  along  inner  orbit, 
gena  along  outer  orbit,  ventral  margin  of 
scape,  maxillary  and  labial  palpi,  prono- 
tum  posteriorly  and  laterally,  discs  of 
scutellum  and  metanotum,  posterior  rim 
of  propodeum,  procoxa,  and  sometimes 
interantennal  tubercle  ventrally,  labrum, 
and  episternum  posteriorly.  Apical  4  or  5 
flagellomeres  black  dorsally.  Basilateral 
and  posterior  portions  of  metasomal  ter- 
gum  I  and  posterior  portions  of  following 
terga  more  or  less  darkened.  Metatarsus 
becoming  darker  towards  apex.  Wings 
hyaline  with  yellowish  brown  tint,  irides- 
cent in  certain  lights,  narrowly  and  weakly 
infuscate  along  outer  margins.  Pterostigma 
dark  brown.  Punctation:  Body  devoid  of 
punctures.  Pubescence  and  setae:  Pubescence 
on  body  usually  very  short  and  decum- 
bent; metanotum,  lateral  portion  of  meta- 
postnotum,  and  propodeum  with  long, 
sub-erect,  brown  pubescence.  Vertex,  la- 


brum, mandible,  propleuron,  and  sterna  V- 
VI  with  short  yellowish  brown  to  brown 
setae;  remainder  of  body  and  legs  almost 
devoid  of  setae.  Head:  1.1-1.2  (1.1  )x  as 
broad  as  long.  Vertex  moderately  to 
strongly  convex  between  eye  tops 
(Fig.  1C).  Frons  distinctly  tuberculate  be- 
tween antennal  sockets  (Figs  IB,  4B);  upper 
frons  broadly  depressed  along  median  line, 
the  latter  being  sharply  impressed  on 
antennal  tubercle  but  becoming  obscure 
near  anterior  ocellus.  Antennocular  line 
depressed  beside  antennal  tubercle 
(Fig.  1A).  Inner  orbits  distinctly  emarginate 
at  upper  1/3,  gently  convergent  below 
(Fig.  1C).  UID:MID:LID  =  9.1-9.4:10:6.3-6.7 
(9.4:10:6.3).  MID  0.57-0.60  (0.58)  X  head 
width.  Ocelli  forming  acute  triangle,  this 
area  being  distinctly  raised.  POL:OOL  = 
1:2.5-3.1  (3.1).  Clypeus  slightly  convex, 
1.8-2.0  (2.0)  X  as  broad  as  long;  anterior 
margin  truncate,  weakly  and  arcuately 
emarginate  (Fig.  1C);  lateral  sides  strongly 
convergent  towards  apex.  Labrum  1.8-2.1 
(2.1)  X  as  broad  as  long;  anterior  margin 
feebly  and  triangularly  emarginate.  Man- 
dible narrowly  rounded  without  sharp 
carina  laterally.  Malar  space  short  (Fig.  IB). 
Genae  0.4-0.5  (0.5) X  eye  width  in  profile, 
roundly  receding  in  dorsal  view.  Scape 
with  carina  long  but  not  sharp  beneath; 
face  slightly  concave  laterally  in  dorsal 
view.  Flagellomere  I  2.1-2.9  (2.5)  X  as  long 
as  wide  and  0.34-0.48  (0.44)  X  as  long  as 
UID;  flagellomeres  I  and  II  in  ratio  of 
10:9.3-11    (10:10).    Mesosoma:    Pronotum 


316  Journal  of  Hymenoptera  Research 

short   (Fig.  1A);   anterior   margin   of  disc  Male. — Very  similar  to  female.   Length: 

arcuately  convex  in  dorsal  view,  its  ante-  Body  7.1-12.4  mm;  fore  wing  7.1-11.4  mm. 

rolateral  corner  rounded;  lower  anterolat-  Head:  1.1-1.2X  as  broad  as  long.  UID:MID: 

eral   tubercle   not   much   swollen,   being  LID  =  9.0-9.3:10:6.5-6.8.    MID   0.57-0.62X 

almost  concealed  by  disc  in  dorsal  view  head  width.  POL:OOL=  1:2.3-2.6.  Clypeus 

(compare  Fig.  1A  with  Fig.  1J:  /.  pernix);  1.8-2.0X   as  broad  as  long.  Labrum  1.8- 

posterior  margin  arcuate  with  small  medi-  1.9  X  as  broad  as  long.  Gena  0.4-0.5  X  eye 

an  notch.  Mesoscutum  with  scutal  groove  width  in  profile.  Flagellomere  I  2.0-2.5  X  as 

sharply   impressed   anteriorly,   becoming  long  as  wide,  0.34-0.43  X  as  long  as  UID; 

shallower   and   broader   posteriorly,   but  flagellomeres  I  and  II  in  ratio  of  10:9.2-11. 

almost   attaining   scuto-scutellar   sulcus;  Mesosoma:  Metapostnotum  0.8-1  X   length 

parapsidal   sulcus   appearing   as   a    fine,  of   metanotum   at   midline.    Legs:   Longer 

raised  line;  posterolateral  margin  broadly  spur   of   metatibia   0.69-0.76 X    length   of 

reflexed.  Discs  of  scutellum  and  metano-  metatarsomere  I.  Wings:  Fore  wing  SMC2 

turn  remarkably  projecting,  the  latter  steep-  receiving  crossvein  lm-cu  at  basal  0.56- 

ly  falling  posteriorly  (Fig.  ID).  Posterome-  0.70.  SMC3  narrowed  on  vein  Rs  by  0.67- 

dian  lobes  of  mesosternum  well  developed  0.79X  its  length  on  vein  M,  1.1-1.4X  length 

and   digitate,  apices  close  to  each   other  of  SMC2  on  vein  M,  1.1-1.4X  as  long  as 

(Fig.  IE).  Metapostnotum  0.7-1  (0.7)  X   as  SMC2  on  vein  Rs,  receiving  crossvein  2m- 

long   as   metanotum   at   midline,   deeply  cu    at   basal    0.55-0.67.    Subgenital   plate 

sunken  between   metanotum  and   propo-  (Fig.  1H):  Lateral  sides  gently  convergent 

deum  (Fig.  ID),  with  fine,  transverse  striae,  towards  apex;  apical  margin  sub-triangu- 

Propodeum  strongly  depressed  along  ante-  larly  convex;  ventral  surface  covered  with 

rior   margin,   almost   linearly   sloping   in  minute  setae  except  for  subapical  portion, 

profile,  with  one  or  two  lateral  tubercles  Genitalia  (Fig.  1G):  Paramere  with  strong 

between     spiracle     and     posterior     rim  setae  apicomedially;  parapenial  lobe  slight- 

(Fig.  1A);   infrastigmal   tubercle   roundly  ly  extending  beyond  apex  of  aedeagus. 

raised;   median   groove   impressed   only  Distribution. — Japan   (Honshu,   Shikoku, 

anteriorly;  surface  smooth,  not  striate.  Me-  and  Kyushu),  Korea,  Taiwan  (Yasumatsu 

tasoma:  Slender  and  almost  parallel-sided  1933),  and  China  (Fig.  6). 
medially.  Tergum  I  abruptly  narrowed,  not 

petiolate   or   parallel-sided   basally.    Legs:  T^e  ^erial.-Holotype  9  (ELKU),  Japan, 

Longer  spur  of  metatibia  0.69-0.77  (0.73)X  K^lshup  M*"   Hik°san'  ^r1**   £.uYasUr 

,                                                T    _        , ,     ,  matsu.  Paratypes:  Japan:  Kyushu:  Mt.  Hikosan, 

as    ong  as  metatarsomere  I.  Tarsal  claws  Fukuoka  Pref     5  viiU940/  K    yasumatsu,  1«J 

with  vertical  tooth  near  middle  (Fig.  IF).  (ELKU).  Lake  Yamashita,  Kokonoe-machi,  Oita 

Wings:   Fore  and   hind   wing  venation   as  Pref.,  9.ix.l997,  R.  Matsumoto,  lo  (OMNH).  Mt. 

shown  in  Fig.  5A.  Fore  wing  crossvein  2r-rs  Ariake-yama,   Izuhara-machi,   Tsushima   Is., 

originating  beyond  middle  of  pterostigma.  24.vii.2001,   R.   Oomuta,    1 J   (TMUB).   Japan: 

Crossvein   cu-a  originating   at  or   slightly  Shikoku:    Mt.    Ishizuchi-san,   Omogo-mura, 

distal  to  point  of  separation  of  vein  M+CuA.  Ehime   Pref.,    17.viii.2002,    M.    Shiraishi,    lo 

SMC2  trapezoid,  receiving  crossvein  lm-cu  (TMUB).  Omogo,  Omogo-mura,  Ehime  Pref., 

at  basal  0.54-0.70  (0.58).  SMC3  narrowed  on  16.viii.1951,  T.  Esaki,  1 J  (ELKU);  23.viii.1953,  T. 

vein  Rs  by  0.70-0.82  (0.77)  X  its  length  on  EdashiSe'  W  (TMUB);  23.ix.1999,  A.  Shimizu,  29 

,,«;,-.  \/f   1  i    1  a  n  qw        i               Cw^n  (TMUB).  Japan:  Honshu:  Jomine  Shrine,  Yano, 

vein  M,  1.1-1.4  (1.3) X  as  long  as  SMC2  on  \,      ..      J .             „  .        J  _,    ,    _    ...   '    .      ' 

\/f    1  t   1  r  /i  a\.,        i               ™  ,^~  Kamnzumi-mura,  Saitama  Pref.,  3.vm.l994,  T. 

vein  M,  1.2-1.5  (1.4)  X  as  long  as  SMC2  on  M      ,       1im;Trm    ~        ,.'                  '    . 

.                    .    .         '                  b  Nambu,   1J   (TMUB).  Onouchi,  Ogano-machi, 

vein  Rs,  receiving  crossvein  2m-cu  at  basal  Saitama  Pref >#  10.x.1992/  T.  Nambll/  1o  (TMUB). 

0.55-0.61  (0.55).  Hind  wing  crossvein  rs-m  Onagata,  Yoshida,  Saitama  Pref.,  18.viii.1988,  T. 

straight,  oblique  to  vein  M.  Crossvein  cu-a  at  Nambu,  69IJ  (TMUB),  19  (FSAG);  26.viii.1988 

ngle  of  approximately  150°  to  vein  A.  (29:  TMUB,  FSAG),  1,  3.viii.l994  (I9:  TMUB),  24, 


Volume  16,  Number  2,  2007 


317 


26.viii.1995  (2Q:  TMUB),  28.vii.2001  (26*:  TMUB), 
A.  Shimizu.  Riv.  Ochi-gawa,  Otaki-mura,  Chi- 
chibu,  Saitama  Pref.,  6.ix.l970,  T.  Nambu,  29 
(TMUB);  6.ix.l999,  A.  Shimizu,  19  (TMUB). 
Kawamata,  Otaki,  Chichibu,  18.viii.2005,  A. 
Shimizu,  I9  (TMUB).  Mt.  Komaga-take, 
1050  m,  Hakone,  Kanagawa  Pref.,  8.viii.2005, 
A.  Shimizu,  I3  (TMUB).  East  of  Fujikawagu- 
chiko-machi,  Minami-tsuru-gun,  Yamanashi 
Pref.,  5.viii.2006,  H.  Takahashi,  26*  (TMUB). 
Mt.    Sanage,    Evergreen    forest,    Aichi    Pref., 


deciduous  forest,  8.ix.2002,  P.  Tripotin,  I9. 
Jeollanamdo,  Gurye-gun,  Toji-myeon,  Nae- 
dong-li,  Piakol  Valley,  on  wild  wine  flowers, 
3.viii.2001,  P.  Tripotin,  19. 

Etymology. — This  species  is  named  after 
the  type  locality. 

Remarks. — This  new  species  is  similar  to 
I.  pernix,  but  the  following  characters 
distinguish  them: 


/.  hikosanus 

1.  Lower  anterolateral  tubercle  of  pronotum  slightly 
and  roundly  produced,  almost  concealed  by  disc 
in  dorsal  view  (Fig.  1A). 

2.  Interantennal  tubercle  merging  into  upper  frons, 
with  median  line  finely  impressed  (Fig.  4B). 


/.  pernix 
Lower  anterolateral  tubercle  of  pronotum  angulate, 

markedly  projecting  beyond  disc  in  dorsal  view 

(Fig.  1J). 
Interantennal  tubercle  abruptly  raised  from  slightly 

depressed  upper  frons,  with  median  line  deeply 

and  broadly  impressed  (Fig.  4A). 


28.viii-3.ix. 1992,  T.  Kanbe,  Malaise  trap,  I9 
(TMUB).  Hio,  Kanazawa-shi,  Ishikawa  Pref., 
27.viii,  1998,  Y.Tazaki,  1 6*  (NSMT).  Misaka-dani, 
Izumi-mura,  Ono-gun,  Fukui  Pref.,  13.ix.2002, 
H.  Takahashi,  I9  (TMUB).  Kaizuka-shi,  Izumi- 
katsuragisan,  Osaka,  l.x.2000,  R.  Matsumoto,  I9 
(OMNH);  13-23.vii.2002  (16*),  23.vii-2.viii.2002 
(26*),  2-10.viii.2002  (23),  20.viii-2.ix.2002  (I9),  2- 
14.ix.2002  (19),  23.ix-2.x.2002  (I9),  2-11.X.2002 
(16*),  Malaise  trap,  R.  Matsumoto,  (OMNH). 
Kishiwada-shi,  Izumi-katsuragisan,  Osaka,  20- 
30.vi.2002  (19),  30.vi-13.vii.2002  (I9I6*),  13- 
23.vii.2002  (29),  23.vii-2.viii.2002  (16*),  2- 
10.viii.2002  (26*),  10-20.viii.2002  (l;),  23.ix-2.x. 
2002  (49),  Malaise  trap,  R.  Matsumoto,  (OMNH). 
Six  stage  of  Mt.  Atago-yama,  Ukyo-ku,  Kyoto-shi, 
27.viii.1987,  A.  Ichikawa,  19  (OMNH).  Hanase 
Pass,  Kyoto-shi,  10.ix.1999,  R.  Matsumoto,  19 
(OMNH).  Mimuro,  Shingo-cho,  Okayama  Pref., 
6.ix.l992,  R.  Matsumoto,  19  (OMNH).  Kozagawa, 
Wakayama  Pref.,  20. ix. 1957,  S.  Momoi,  39 
(TMUB).  Daisen,  Tottori  Pref.  (Hoki),  19.viii.1932, 
S.  Yasimoto,  1J  (ELKU).  Korea:  Chungcheong- 
namdo,  Keumsan,  Poseoksa,  10.viii.1998  (I9), 
22.viii.1998  (I9),  24.ix.2000  (I9),  ix.2001  (29),  P. 
Tripotin,  (FSAG).  Kyeongsangnamdo,  Jirisan, 
Hamyang-gun,  Macheon-myon,  Samjeong-li  Jir- 
isan, 700  m,  23-25.viii.2002  (49),  10-20.ix.2003 
(16*),  Malaise  trap,  P.  Tripotin,  (FSAG).  CHINA: 
Szechwan,  Suifu,  1000-1500  m,  l-21.vi.1928,  D. 
Graham,  19  (USNM). 

Non-type    material. — Korea:    Chungcheong- 
namdo,    Keumsan,    Poseoksa,    along    trail    in 


Irenangelns  nambui  Shimizu,  new  species 

(Hg.  2) 

Female. — Length:  Body  4.5-7.6  (5.5)  mm; 
fore  wing  4.7-6.9  (5.2)  mm.  Coloration: 
Body  predominantly  blackish  brown  to 
black  and  polished.  Following  ivory-white: 
clypeus  and  labrum  laterally,  mandible 
(apical  portion  brown),  ventral  margin  of 
scape,  maxillary  and  labial  palpi,  procoxa 
(basal  portion  more  or  less  dark  brown), 
protrochanter,  and  sometimes  profemur, 
mid  and  hind  coxae,  trochanters,  femora 
and  tibiae  partly.  Remainder  of  fore  leg 
light  brown.  Mid  and  hind  legs  predomi- 
nantly brown,  darker  than  fore  leg,  but 
somewhat  lighter  ventrally  than  dorsally. 
All  tibial  spurs  ivory-white  to  yellowish 
light  brown.  Posterolateral  margin  of  pro- 
notum, lateral  and  posterior  portions  of 
metasomal  terga,  and  posterior  portions  of 
metasomal  sterna  light  brown  to  ferrugi- 
nous. Wings  hyaline,  iridescent  in  certain 
lights,  weakly  infuscate  along  outer  mar- 
gins. Pterostigma  dark  brown.  Punctation: 
Body  devoid  of  punctures.  Pubescence  ami 
setae:  Body  and  legs  with  short,  appressed 
white  pubescence,  longer  and  denser  on 
lower  frons,  clypeus,  lower  pronotum, 
propleuron,  mesopleuron,  lower  meta- 
pleuron,   propodeum,   and   coxae.    Upper 


318 


Journal  of  Hymenoptera  Research 


Fig.  2.  Irenangelus  nambui  n.  sp.  (A-G,  holotype  female;  H-I,  paratype  male  from  Japan).  A,  Whole  body, 
dorsal  view;  B,  head,  frontal  view;  C,  head,  pronotum,  and  mesoscutum,  dorsal  view;  D,  head,  lateral  view.  E, 
mesosternum  and  mesocoxa,  ventral  view;  F,  mesosoma,  lateral  view;  G,  right  metatarsal  claw,  outer  view;  H, 
subgenital  plate,  ventral  view.  I,  genitalia  (left  half,  ventral  view;  right  half,  dorsal  view).  Scale  lines:  0.5  mm. 


irons,  vertex,  clypeus,  labrum,  mandible, 
apices  of  terga  VI  and  sterna  IV-V,  and 
sternum  VI  with  short  pale  setae.  Head: 
1.2X  as  broad  as  long.  Vertex  strongly 
convex  in  frontal  view  (Fig.  2B).  Frons  with 
interantennal  area  not  tuberculate  but 
slightly  overhanging  antennal  sockets 
(Fig.  2D);  median  line  impressed  only  on 
lower  half.  Antennocular  line  nearly  trans- 
verse (Fig.  2C).  Inner  orbits  slightly  emar- 
ginate  a  little  above  middle,  strongly  di- 
vergent above.  UID:MID:LID=9.8-10.1:10: 
6.8-7.1  (10:10:6.9).  MID  0.57-0.60  (0.60) X 
head  width.  Ocelli  forming  acute  triangle, 
this  area  being  scarcely  raised.  POL: 
OOL=  1:2.5-3.6  (1:3.6).  Clypeus  slightly 
convex,  2.1-2.4  (2.2)  X  as  broad  as  long; 
anterior  margin  truncate,  weakly  and 
arcuately  emarginate  (Fig.  2B);  lateral  sides 
arcuately   convergent   towards   apex.   La- 


brum 2.5-3.0  (3.0)  X  as  broad  as  long; 
anterior  margin  arcuately  emarginate. 
Mandible  carinate  laterally.  Malar  space 
short.  Genae  0.3-0.4  (0.4)  X  eye  width  in 
profile,  roundly  receding  in  dorsal  view. 
Scape  sharply  carinate  beneath;  lateral  face 
slightly  concave  in  dorsal  view.  Flagello- 
mere  I  2.9-3.5  (3.0)  X  as  long  as  wide  and 
0.45-0.49  (0.47)  X  length  of  UID;  flagello- 
meres  I  and  II  in  ratio  of  1:0.94-1.0  (1:0.98). 
Mesosoma:  Pronotum  short;  anterior  margin 
of  disc  arcuately  convex  in  dorsal  view,  its 
anterolateral  corner  gently  rounded 
(Fig.  2C);  lower  anterolateral  tubercle  not 
much  swollen,  being  completely  concealed 
by  disc  in  dorsal  view;  posterior  margin 
arcuate  with  small  median  notch.  Mesos- 
cutum with  scutal  groove  sharply  im- 
pressed anteriorly,  becoming  shallower 
and  broader  posteriorly,  obsolete  just  an- 


Volume  16,  Number  2,  2007 


319 


terior  to  scuto-scutellar  sulcus;  parapsidal 
sulcus  finely  impressed;  posterolateral 
margin  narrowly  reflexed.  Discs  of  scutel- 
lum  and  metanotum  distinctly  projecting 
(Fig.  2F),  the  latter  being  pyramidal.  Pos- 
teromedian lobes  of  mesosternum  short 
but  bilobed,  their  inner  lobes  close  to  each 
other  (Fig.  2E).  Metapostnotum  0.7-0.9 
(0.7)  X  length  of  metanotum  at  midline, 
with  few  very  fine  striae  anteriorly  and 
distinct  longitudinal  median  groove.  Pro- 
podeum  weakly  convex  in  profile  (Fig.  2F), 
scarcely  depressed  along  anterior  margin, 
without  lateral  tubercle  or  infrastigmal 
tubercle  (Fig.  2A);  median  groove  obsolete; 
surface  smooth,  not  striate.  Metasoma: 
Slenderly  fusiform.  Tergum  I  abruptly 
narrowed,  not  petiolate  or  parallel-sided 
basally.  Legs:  Longer  spur  of  metatibia 
0.65-0.74  (0.69)  X  metatarsomere  I.  Tarsal 
claws  sub-bifid:  inner  ray  sub-parallel  to 
outer  ray,  acute.  Wings:  Fore  and  hind 
wing  venation  as  shown  in  Fig.  2A.  Fore 
wing  crossvein  2r-rs  originating  before 
middle  of  pterostigma.  Crossvein  cu-a  orig- 
inating slightly  basad  of  point  of  separation 
of  vein  M+CuA.  SMC2  rhomboid,  receiving 
crossvein  lm-cu  at  basal  0.43-0.56  (0.52). 
SMC3  narrowed  on  vein  Rs  by  0.42-0.53 
(0.51)  X  its  length  on  vein  M,  1.2-1.5  (1.3)  X 
as  long  as  SMC2  on  vein  M,  0.69-1.1  (0.87)  X 
as  long  as  SMC2  on  vein  Rs,  receiving 
crossvein  2m-cu  at  apical  0.54-0.66  (0.58). 
Hind  wing  crossvein  rs-m  almost  straight, 
oblique  to  vein  M.  Crossvein  cu-a  forming 
angle  of  135-140   to  vein  A. 

Male. — Very  similar  to  female.  Length: 
Body  3.9-8.3  mm;  fore  wing  3.7-6.5  mm. 
Head:  1.2X  as  broad  as  long.  UID:MID: 
LID=9.9-10.1:10:7.0-7.7.  MID  0.57-0.60  X 
head  width.  POL:OOL=l:2.1-3.1.  Clypeus 
2.3-2.5  X  as  broad  as  long.  Labrum  2.6- 
3.2  X  as  broad  as  long.  Gena  0.3-0.4  X  eye 
width  in  profile.  Flagellomere  I  2.2-2.7  X  as 
long  as  wide  and  0.38-0.43 X  as  long  as 
UID;  flagellomeres  I  and  II  in  ratio  of 
1:0.96-1.1.  Mesosoma:  Metapostnotum  0.8- 
lx  length  of  metanotum  at  midline.  Legs: 
Longer  spur  of  metatibia  0.66-0.73 X  meta- 


tarsomere I.  Wings:  SMC2  receiving  cross- 
vein  lm-cu  at  basal  0.44-0.60.  SMC3 
narrowed  on  vein  Rs  by  0.44-0.73  X  its 
length  on  vein  M,  1.1-1.6X  as  long  as 
SMC2  on  vein  M,  0.64-1 .4  X  as  long  as 
SMC2  on  vein  Rs,  receiving  crossvein  2m- 
cu  at  apical  0.50-0.66.  Subgenital  plate 
(Fig.  2H):  Broadened  medially;  apical  mar- 
gin slightly  emarginate  or  truncate;  ventral 
surface  covered  with  minute  setae.  Genita- 
lia (Fig.  21):  Paramere  without  strong  setae 
apicomedially;  parapenial  lobe  short,  not 
attaining  apex  of  aedeagus. 

Distribution. — Japan  and  Korea  (Fig.  6). 

Type  material. — Holotype  9  (TMUB),  Nageishi 
Pass,  Higashi-Mikabo,  Gunma  Prefecture, 
28.viii.1986,  T.  Nambu.  Paratypes:  Japan:  Hon- 
shu: Showa,  Mt.  Hakase,  1000  m,  Beech  forest, 
Fukushima  Pref.,  29.vi-26.vii.1998  (1  ;),  27.vii- 
23.viii.1998  (697^),  24.viii-19.ix.1998  (192^),  Ma- 
laise trap,  T.  Muroi,  (TMUB).  Imperial  Palace, 
Chiyoda-ku,  Tokyo,  28.V.1999,  T.  Nambu,  1<$ 
(TMUB).  Mt.  Komaga-take,  1000-1300  m,  Ha- 
kone,  Kanagawa  Pref.,  ll.vii.2000  (1^), 
30.viii.2000  (I9),  H.  Nagase,  (TMUB);  18.vii.2001 
(1976*:  TMUB;  1  £  FSAG),  l.viii.2001  (26*:  TMUB), 
A.  Shimizu.  Takekurabe-yama,  Maruoka-cho, 
Fukui  Pref.,  5.ix.l994,  Y.  Haneda,  I9  (TMUB). 
Akausagi-yama,  Ohno-shi,  23. ix. 1974,  Y.  Ha- 
neda, 19  (FSAG).  Shitara,  Beech  forest,  900  m, 
Uradani,  Aichi  Pref.,  25-31.vii.1994  (1  J),  29.viii- 
4.ix.l994  (1  ;),  Malaise  trap,  K.  Yamagishi, 
(TMUB);  29.viii-4.ix.1994,  Emergence  trap,  K. 
Yamagishi,  I9I 5  (TMUB);  19-25.ix.1994,  Pan 
trap,  K.  Yamagishi,  I9  (TMUB);  l-7.viii.1994 
(1J:  TMUB),  22-28.viii.1994  (I9I6*:  TMUB;  19, 
FSAG),  Malaise  trap,  T.  Kanbe.  Asahi,  Yawata, 
650  m,  Deciduous  forest,  17-26.vi.1998  (2;),  12- 
21.viii.1998  (I9I ?),  15-25.ix.1998  (19),  Malaise 
trap,  M.  Ozawa,  (TMUB).  Mt.  Sanage,  Evergreen 
forest,  Aichi  Pref.,  28.viii-3.ix.1992,  Emergence 
trap,  K.  Shima,  I9  (TMUB);  4-10.ix.1992,  Malaise 
trap,  T.  Kanbe,  19  (TMUB);  16-22.ix.2002,  Ma- 
laise trap,  M.  Kiyota,  1  J  (TMUB).  Korea: 
Kyeongsangnamdo,  Jirisan,  Hamyang-gun,  Ma- 
cheon-myon,  Samjeong-li,  700  m,  35  20'55N 
127  38'21E,  Malaise  trap,  10-20.ix.2003,  P.  Tripo- 
tin,  292;  (FSAG). 

Etymology. — This  species  is  named  in 
honor  of  the  provider  of  the  holotype 
specimen. 


320 


Journal  of  Hymenoptera  Research 


Fig.  3.  Irenangelus  punctipleuris  n.  sp.  (A-G,  I,  holotype  female;  H,  J-K,  paratype  females:  H,  from  Sulawesi;  J, 
from  Brunei;  K,  from  Japan);  L-M,  paratype  male  from  Japan).  A,  Head  and  antenna,  lateral  view;  B,  head, 
frontal  view;  C,  head,  dorsal  view;  D,  mesosoma,  lateral  view;  E,  metasomal  tergum  I,  dorsal  view;  F, 
mesosternum  and  mesocoxa,  ventral  view;  G,  right  metatarsal  claw,  outer  view;  H-K,  pronotum,  dorsal  view;  L, 
genitalia  (left  half,  dorsal  view;  right  half,  ventral  view);  M,  sternum  VII  and  subgenital  plate,  ventral  view.  Scale 
lines:  0.5  mm. 


Remarks. — In  Irenangelus  this  species  is 
unique  in  its  predominantly  dark  brown  to 
black  body  and  wholly  transparent  wings. 

Irenangelus  punctipleuris  Wahis,  new 
species 

(Figs  3,  4C-F,  5B-C) 

Female. —Length:  Body  8.1-12.4  (10.0) 
mm;  fore  wing  6.2-9.3  (7.9)  mm.  Coloration: 
Head,  mesosoma  and  coxae  black  with 
following  bright  yellow:  clypeus  and  la- 
brum  (lateral  portions  black),  frons  be- 
tween and  below  antennal  sockets,  upper 
frons  along  inner  orbit  and   gena   along 


outer  orbit  broadly,  scape  and  pedicel 
(dorsal  faces  dark  brown  to  black),  prono- 
tal  disc  (lateral  margin  black),  ventral  and 
posterolateral  margins  of  pronotum  broad- 
ly, posteromedian  elliptic  spot  and  lateral 
streak  on  mesoscutum,  median  spot  on 
scutellum,  metanotal  disc,  oblong  spot  on 
upper  mesopleuron,  this  spot  being  some- 
times obsolete,  two  large  spots  on  lower 
mesopleuron,  these  often  being  continuous 
(Figs  4E-F),  median  triangular  and  lateral 
longitudinal  marks  on  propodeum,  these 
being  continuous  posteriorly,  oblique  spot 
on   upper   metapleuron,    this   spot   being 


*  .      a  ■     - 


Volume  16,  Number  2,  2007 


321 


Fig.  4.  Female  head  (A-D)  and  mesopleuron  (E-F)  of  Irenangelus  (A-B,  dorsolateral  view;  C-D,  frontal  view; 
E-F,  lateral  view).  A,  /.  pernix  Bingham  from  Bali,  Indonesia;  B,  /.  hikosanus  n.  sp.,  holotype;  C-F,  /.  punctipleuris, 
n.  sp.  (C,  E,  holotype;  D,  F,  paratype  from  Japan). 


sometimes  obsolete,  posterodorsal  mark  on 
lower  metapleuron,  this  being  continuous 
with  lateral  propodeal  mark,  apical  greater 
parts  of  coxae,  and  sometimes  side  of 
metanotum  and  anterior  portion  of  meta- 
postnotum.  Following  reddish  brown:  fla- 
gellomeres  I-IV  or  -X  ventrally  (remainder 
of  flagellum  dark  brown  to  black),  maxil- 


lary and  labial  palpi,  trochanters  (basally 
dark  brown  to  black),  femora  (dorsolateral 
portion  of  profemur  and  sometimes  ventral 
portions  of  meso-  and  metafemora  bright 
yellow;  sometimes  all  femora  dark  brown 
ventrally  and/or  laterally),  tibiae  (protibia 
bright  yellow  dorsolaterally;  sometimes 
dorsal  portion  of  mesotibia  and  basidorsal 


Fig.  5.     Female  wings  (A,  fore  and  hind  wings;  B,  fore  wing;  C,  hind  wing).  A,  /.  hikosanus  n.  sp.,  holotype;  B-C, 
/.  punctipleuris  n.  sp.,  paratype  from  the  Philippines. 


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portion  of  metatibia  dark  brown  to  black), 
tarsi,  and  metasoma  (tergum  I  dark  brown 
to  black  anterodorsally  and  sublaterally; 
terga  II— IV  sometimes  becoming  darker 
dorsally).  Posterolateral  margin  of  mesos- 
cutum,  tegula,  and  wing  bases  yellowish 
brown.  Mandible  black;  apical  1/3  dark 
rufous.  Wings  hyaline,  iridescent  in  certain 
lights,  weakly  infuscate  along  outer  mar- 
gins. Pterostigma  light  to  dark  brown. 
Punctation:  Upper  frons  (Figs  4C-D),  vertex 
between  eye  and  ocellus,  pronotum,  me- 
soscutum,  discs  of  scutellum  and  metano- 
tum,  mesopleuron  (Figs  4E-F),  and  lower 
metapleuron  with  irregularly-spaced,  shal- 
low punctures,  these  being  larger  and 
denser  along  median  line  of  frons  and 
scutal  groove,  and  sometimes  on  meso- 
pleuron. Pubescence  and  setae:  Body  and  legs 
with  short,  appressed,  white  to  pale  brown 
pubescence  but  devoid  of  long  bristly 
setae;  vertex,  mandible,  propleuron,  lateral 
side  of  pronotal  disc,  mesopleuron,  meta- 
notum,  and  posterolateral  portion  of  pro- 
podeum  with  short,  dense,  white  to  pale 
brown  setae.  Head:  1.1-1.2  (1.2)X  as  broad 
as  long.  Vertex  strongly  convex  in  frontal 
view  (Fig.  3B).  Frons  without  interantennal 
tubercle  (Fig.  3A);  median  line  finely  im- 
pressed from  interantennal  area  close  to 
anterior  ocellus.  Antennocular  line  slightly 
depressed  nearby  antennal  base  (Fig.  3C). 
Inner  orbits  distinctly  emarginate  at  upper 
1/3,  strongly  convergent  below.  UID: 
MID:LID  =  8. 5-9.4:10:5.6-6.5  (9.3:10:5.9). 
MID  0.59-0.63  (0.61)  x  head  width.  Ocelli 
forming  slightly  acute  triangle,  this  area 
being  slightly  raised.  POL:OOL  =  1:1. 6-2.5 
(1.9).  Clypeus  feebly  convex,  1.6-1.9  (1.8) X 
as  broad  as  long;  anterior  margin  truncate, 
weakly  and  arcuately  emarginate  (Fig.  3B); 
lateral  sides  strongly  convergent  towards 
apex.  Labrum  2.0-2.3  (2.2)  X  as  broad  as 
long;  anterior  margin  with  small  median 


notch.  Malar  space  very  short  (Fig.  3A). 
Mandible  sharply  carinate  laterally.  Gena 
0.4-0.5  (0.4)  X  eye  width  in  profile,  feebly 
rounded  in  dorsal  view.  Scape  carinate  on 
apical  half  ventrally;  lateral  face  flattened 
but  scarcely  concave  in  dorsal  view.  Fla- 
gellomere  I  2.1-2.4  (2.4)  X  as  long  as  wide 
and  0.38-0.43  (0.38)  X  length  of  UID; 
flagellomeres  I  and  II  in  ratio  of  1:0.84-1.0 
(0.94).  Mesosoma:  Pronotum  short;  anterior 
margin  of  disc  almost  straight  in  dorsal 
view,  its  lateral  corner  sub-angulate,  but 
degree  of  angulation  variable  (Figs  3H-K); 
lower  anterolateral  tubercle  not  much 
swollen,  being  concealed  by  disc  in  dorsal 
view;  posterior  portion  narrowly  but  dis- 
tinctly depressed  along  posterior  margin, 
the  latter  being  arcuate.  Mesoscutum  with 
scutal  groove  shallowly  impressed  on 
anterior  1/4-3/4;  parapsidal  sulcus  ap- 
pearing as  a  fine,  raised  line;  posterolateral 
margin  narrowly  reflexed.  Discs  of  scutel- 
lum and  metanotum  strongly  raised 
(Fig.  3D).  Metapostnotum  0.59-1.0  (1.0) X 
length  of  metanotum  at  midline,  deeply 
sunken  between  metanotum  and  propo- 
deum,  with  fine,  transverse  striae  anterior- 
ly and  short  oblique  striae  posteriorly. 
Posteromedian  lobes  of  mesosternum  tri- 
angularly produced,  their  apices  removed 
from  each  other  (Fig.  3F).  Upper  meta- 
pleuron finely  and  obliquely  striate.  Pro- 
podeum  short,  barely  convex  in  profile 
(Fig.  3D),  deeply  depressed  along  anterior 
margin,  without  lateral  tubercle;  infrastig- 
mal  tubercle  weak;  surface  finely  and 
transversely  striate,  with  weak  to  rudimen- 
tary median  groove.  Metasoma:  Much  slen- 
derer than  mesosoma.  Tergum  I  gradually 
narrowed  and  petiolate  or  parallel-sided 
basally  (Fig.  3E).  Legs:  Longer  spur  of 
metatibia  0.63-0.72  (0.68)  X  length  of  meta- 
tarsomere  I.  Tarsal  claws  bifid;  inner  ray  of 
claw  truncate  (Fig.  3G).   Wings:   Fore  and 


Fig.  6.     Map  showing  the  known  distribution  of  Irenangelus  hikosanus,  I.  nambui,  and  /.  punctipleuris.  In  fapan 
only  certain  localities  of  specimens,  including  type  localities,  have  been  selected  for  /.  hikosanus  and  /.  nambui. 


324 


Journal  of  Hymenoptera  Research 


hind  wing  venation  as  shown  in  Figs  5B  and 
C,  respectively.  Fore  wing  crossvein  2r-rs 
originating  slightly  before  middle  of  pter- 
ostigma.  Crossvein  cu-a  usually  originating 
at  or  slightly  basad  of  point  of  separation  of 
vein  M+CuA.  SMC2  almost  rectangular, 
receiving  crossvein  lm-cu  at  basal  0.43- 
0.57  (0.54).  SMC3  narrowed  on  vein  Rs  by 
0.75-0.85  (0.84)  X  its  length  on  vein  M,  1.3- 
1.8  (1.8)x  as  long  as  SMC2  on  vein  M,  1.2- 
1.8  (1.8)  X  as  long  as  SMC2  on  vein  Rs, 
receiving  crossvein  2m-cu  at  basal  0.39-0.61 
(0.50).  Hind  wing  crossvein  rs-m  almost 
vertical  to  vein  M.  Crossvein  cu-a  forming 
angle  of  approximately  150   to  vein  A. 

Male. — Very  similar  to  female.  Length: 
Body  6.7-8.4  mm;  fore  wing  5.7-7.0  mm. 
Head:  1.2X  as  broad  as  long.  UID:MID: 
LID=9.0:10:6.7-7.3.  MID  0.62-0.65X  head 
width.  POL:OOL=l:1.9-2.1.  Clypeus  2.0x 
as  broad  as  long.  Labrum  2. 1-2.2 X  as 
broad  as  long.  Gena  0.4-0.5 X  eye  width 
in  profile.  Flagellomere  I  2.3-2.4  X  as  long 
as  wide  and  0.32-0.35  X  UID;  flagellomeres 
I  and  II  in  ratio  of  1:0.87-0.90.  Mesosoma: 
Metapostnotum  0.75  X  length  of  metano- 
turrt  at  midline.  Legs:  Longer  spur  of 
metatibia  0.72 X  metatarsomere  I.  Wings: 
SMC2  receiving  crossvein  lm-cu  at  basal 
0.42-0.55.  SMC3  narrowed  on  vein  Rs  by 
0.77-0.78  X  its  length  on  vein  M,  1.4-1.7X 
as  long  as  SMC2  on  vein  M,  1.3-1.5  X  as 
long  as  SMC2  on  vein  Rs,  receiving  cross- 
vein  2m-cu  at  basal  0.45-0.67.  Subgenital 
plate  (Fig.  3M):  Lateral  sides  gradually 
convergent  towards  apex;  apical  margin 
sub-triangularly  produced;  ventral  surface 
with  minute  setae  apically.  Genitalia 
(Fig.  3L):  Paramere  with  strong  setae  api- 
comedially;  parapenial  lobe  long  and  slen- 
der, decurved  apically,  extending  beyond 
apex  of  aedeagus. 

Distribution. — From  Japan  through  the 
Philippines  and  Malaysia  to  India  and  Sri 
Lanka  (Fig.  6). 

Type  material— Holotype  9  (ZMUC),  Philip- 
pines, Balabac  Dalawan  Bay,  7.x. 1961,  Noona 
Dan  Exp.  61-62.  Paratypes:  Philippines:  Tawi 
Tawi,  Tarakawan,  north  of  Batu  Batu,  4.xi.l961 


(19:  ZMUC),  10.xi.1961   (19:  FSAG),  12.xi.1961 
(19:  ZMUC),  Noona  Dan  Exp.  61-62.  Brunei: 
Ulu    Temburong,    Base    camp    hut,    300  m, 
115  16'E  4  26'N,  16.ii-9.iii.1982,  M.  C.  Day,  29 
(BMNH,   FSAG).   Sulawesi:   Central  Sulawasi, 
Napu-valley,    100  km   S/O,    Palu,   near    Lore- 
Lindu  National  Park,  9.U.2001,  A.  -M.  Klein,  I9 
(FSAG).  Utara,  Dumoga-Bone  Nat.  Park,  ii.1985, 
19  (BMNH).  Bali:  W.  Bali,  near  Negara,  rain- 
forest above  Batuagung,  550  m,  4-6.xii.1911,  C. 
v.    Achterberg,    I9    (RMNH).    Java:    W.    Java, 
Djampang-Tengah,  Mrs.   Walsh,   19  (RMNH). 
Borneo:  Sarawak,  S.W.  Gunung  Buda,  64  km  S. 
Limbang  4  13'N  114  56'E,  8-15.xi.1996,  Malaise 
trap,  S.   L.   Heydon  &  S.   Fung,   19  (UCDC). 
Malaysia:  S.  E.  Sabah,  Danum  Valley  Field  C, 
117  48'E   4  58'N,   x-xii.1986,   P.   Eggleton,   49 
(BMNH),  19  (FSAG).  S.  E.  Sabah,  near  Danum 
Valley    Field,    ca.150  m,    26.v-20.vi. 1987    (49: 
RMNH;    39:     FSAG),    20.vi-12.vii.1987    (I9: 
RMNH),   13.ix-4.x.l987  (I9:   RMNH),  Malaise 
trap,  C.  v.  Achterberg  &  D.  Kennedy.  S.  W. 
Sabah,  near  Long  Pa  Sia  (West),  1020  m,  25.xi- 
9.xii.l987,  Malaise  trap,  C.  v.  Achterberg,  19 
(RMNH).  S.  W.  Sabah,  near  Long  Pa  Sia  (East), 
1000  m,  l-13.iv.1987  (I9),  25.xi-9.xii.1987  (19), 
Malaise  trap,  C.  v.  Achterberg,  (RMNH).  Pasoh 
Forest  Reserve,  Negeri  S.,  22.vii.1978  (I9:  AEIC), 
7.ix.l978    (19:    FSAG),   6.xi.l978    (I9:    AEIC), 
8.i.l979  (19:  AEIC),  P.  &  M.  Becker.  Bukit  Kutu, 
304.1930,  H.  T.  Padgen,  19  (BMNH).  India:  U.  P. 
Garjia,  610  m,  26-29.iv.1969,  Gupta,  No.  335, 1J 
(FSAG).  S.  India,  Madras  ST.,  Anamalai  Hills, 
3500  f,  v.1964,  P.  S.  Nathan,  19  (CNC).  Kerala, 
Periyar  A.  Sanctuary,  5-15.X.1979,  19  (BMNH). 
Sri  Lanka:  Kandy  District,  Udawattakele  Sanc- 
tuary, 1800  f,  l-3.ix.1980,  Malaise  trap,  K.  V. 
Krombein  et  al,  19  (FSAG).  Japan:  Kawamata, 
Otaki-mura,    Chichibu,    Saitama    Pref.,    30. vi- 
l.vii.2004,   A.   Shimizu,   19  (TMUB).   Maruno- 
machi,    Nirasaki-shi,   Yamanashi    Pref.,    27.vi- 
5.vii.2005,  Malaise  trap,  K.  Hosoda,  13  (TMUB). 
Kanegasaki-cho,    Tsuruga-shi,    Fukui    Pref., 
3.vii.2001,  H.  Takahashi,  19  (TMUB). 

Etymology. — The  species  name  is  derived 
from  the  punctate  mesopleuron;  puncti- 
(punctate)  +  pleuris  (pleuron). 

Remarks. — The  present  species  is  similar 
to  "Ceropales"  tennatus  Turner  1910  occur- 
ring in  Australia  in  that  1)  the  flagellum  is 
crenulate;  2)  all  tarsal  claws  are  bifid;  3)  the 
frons  is  devoid  of  an  interantennal  tubercle; 


Volume  16,  Number  2,  2007 


325 


and  4)  the  metasomal  tergum  I  is  gradually 
narrowed  and  petiolate  or  parallel-sided 
basally.  However,  this  new  species  is  easily 
distinguished  from  the  latter  by  the  almost 
entirely  rufous  metasoma  and  the  distinctly 
punctate  mesopleuron. 

ACKNOWLEDGMENTS 

We  thank  Emeritus  Professor  R.  Ishikawa  (Tokyo 
Metropolitan  University)  and  Dr  N.  Springate  (Natu- 
ral History  Museum,  London)  for  reviewing  our 
manuscript  and  providing  critical  comments.  For  the 
gift  or  loan  of  specimens,  our  thanks  are  also  due  to 
the  following:  M.  C.  Day,  T.  Edashige,  T.  Esaki,  Y. 
Haneda,  K.  Hosoda,  L.  S.  Kimsey  (UCDC),  K.  V. 
Krombein  (USNM),  R.  Matsumoto  (OMNH),  S.  Mo- 
moi,  H.  Nagase,  T.  Nambu,  R.  Oomuta,  A.  Shinohara 
(NSMT),  M.  Shiraishi,  O.  Tadauchi  (ELKU),  H. 
Takahashi,  Y.  Tazaki,  P.  Tripotin,  L.  B.  Vilhelmsen 
(ZMUC),  K.  Yamagishi,  Y.  Yasimoto,  and  K.  Yasu- 
matsu. 

LITERATURE  CITED 

Banks,  N.  1934.  The  Psammocharidae  of  the  Philip- 
pines. Proceedings  of  the  American  Academy  of  Arts 
and  Sciences  69:  1-117. 

Bingham,  C.  T.  1896.  On  some  exotic  fossorial 
Hymenoptera  in  the  collection  of  the  British 
Museum,  with  descriptions  of  new  species  and 
of  a  new  genus  of  the  Pompilidae.  journal  of  the 
Linnean  Society  of  London  (Zoology)  25:  422-A45. 

Cameron,  P.  1891.  Hymenoptera  Orientalis;  or  Con- 
tributions to  a  knowledge  of  the  Hymenoptera  of 
the  Oriental  Zoological  Region.  Part  3.  Memoirs 
and  Proceedings  of  the  Manchester  Literary  and 
Philosophical  Society  (4)  4:  431-481,  pi.  3. 

.  1896.  Hymenoptera  Orientalis;  or  Contribu- 
tions to  a  knowledge  of  the  Hymenoptera  of  the 
Oriental  Zoological  Region.  Part  5.  Memoirs  and 
Proceedings  of  the  Manchester  Literary  and  Philo- 
sophical Society  41  (4):  1-144,  pis  3,  4. 

Day,  M.  C.  1988.  Spider  wasps,  Hymenoptera: 
Pompilidae.  Handbooks  for  the  Identification  of 
British  Insects  6  (4):  1-60. 

Evans,  H.  E.  1969.  Studies  on  Neotropical  Pompilidae 
(Hymenoptera)  VII.  Irenangeliis  Schulz.  Studio 
Entomologica  12:  417-431. 

.  1987.  A  new  species  of  Irenangeliis  from  Costa 

Rica  (Hymenoptera:  Pompilidae:  Ceropalinae). 
Proceedings  of  the  Entomological  Society  of  Washing- 
ton 89:  559-561. 

Kim,  C.-W.  1970.  Illustrated  Encyclopedia  of  Fauna  and 
Flora  of  Korea.  Vol.  11,  part  3.  Samwha-Chulpansa, 
Seoul.  891  pp.  [In  Korean.] 


Kimsey,  L.  S.  and  M.  S.  Wasbauer.  2004.  Revision 
of  New  World  species  of  the  cleptoparasitic 
pompilid  genus  Irenangeliis  Schulz  (Hymenop- 
tera: Pompilidae).  journal  of  the  Kansas  Entomolog- 
ical Society  77:  650-668. 

Lelej,  A.  S.,  T.  Saigusa,  and  C.  E.  Lee.  1994.  Spider 
wasps  (Hymenoptera,  Pompilidae)  of  Korea. 
Russian  Entomological  journal  3:  135-148. 

,   T.   Tano,   and    H.    Kurokavva.    1995.   Spider 

wasps  (Hymenoptera,  Pompilidae)  from  Cheju- 
Do  Island,  Southern  Korea.  Transactions  of  Essa 
Entomological  Society  75:  44-47. 

Pitts,  J.  P.,  M.  S.  Wasbauer,  and  C.  D.  Dohlen.  2006. 
Preliminary  morphological  analysis  of  relation- 
ships between  the  spider  wasp  subfamilies 
(Hymenoptera:  Pompilidae):  revising  an  old 
problem.  Zoologica  Scripta  35:  63-84. 

Rohwer,  S.  A.  1919.  Philippine  wasp  studies.  Part.  1. 
Descriptions  of  new  species.  Bulletin.  Hawaiian 
Sugar  Planters'  Association  Experiment  Station, 
Entomological  Series  14:  5-18. 

Schulz,  W.  A.  1906.  Spolia  Hymenopterologica.  A.  Pape, 
Paderborn.  355  pp. 

Shimizu,  A.  1994.  Phylogeny  and  classification  of  the 
family  Pompilidae  (Hymenoptera).  Tokyo  Metro- 
politan University  Bulletin  of  Natural  History  2: 
1-142. 

.  1996.  Key  to  the  genera  of  the  Pompilidae 

occurring  in  Japan  north  of  the  Ryukyus  (Hyme- 
noptera) (Part  2).  Japanese  Journal  of  Entomology  64: 
496-513. 
-,  M.  S.  Wasbauer,  and  M.  Ujiie.  1998.  Taxo- 


nomic  importance  of  the  female  apicoventral 
setae  on  metatarsomere  V  of  the  Pompilidae 
(Hymenoptera),  with  special  reference  to  phvlog- 
eny  of  the  family.  Entomological  Science  1:  427-439. 

Turner,  R.  E.  1910.  Additions  to  our  knowledge  of  the 
fossorial  wasps  of  Australia.  Proceedings  of  the 
Zoological  Society  of  London  1910:  253-356,  pis  31,  32. 

Wahis,  R.  1988.  Hymenopteres  Pompilides  de  Mada- 
gascar. Genus  Ceropales  Latreille  et  Irenangeliis 
Schulz.  (Hymenoptera:  Pompilidae).  Revue  de 
Zoologie  Africaine  102:  213-221. 

Wcislo,  W.  T.,  M.  J.  West-Eberhard,  and  W.  G. 
Eberhard.  1988.  Natural  history  and  behavior  of 
a  primitively  social  wasp,  Auplopus  semialatus,  and 
its  parasite,  Irenangeliis  eberhardi  (Hymenoptera: 
Pompilidae).  Journal  of  Insect  Behavior  1:  247-260. 

Williams,  F.  X.  1919.  Philippine  wasp  studies.  Part  2. 
Descriptions   of   new   species   and    life    history 
studies.  Bulletin.  Hawaiian  Sugar  Planters'  Associ 
at  ion  Experiment  Station,  Entomological  Series   14: 
19-186. 

Yasumatsu,  K.  1933.  [Two  unrecorded  Psammochar- 
idae from  Japan  and  Formosa.]  Fukuoka  Hakubu- 
tsugaku  Zasshi,  1:  143-149.  [In  Japanese.] 


J.  HYM.  RES. 
Vol.  16(2),  2007,  pp.  326-335 

Natural  History  and  Larval  Behavior  of  the  parasitoid  Zatypota  petronae 

(Hymenoptera:  Ichneumonidae) 

Ju-Lin  Weng*  and  Gilbert  Barrantes1 

Escuela  de  Biologia,  Universidad  de  Costa  Rica,  Ciudad  Universitaria  Rodrigo  Facio,  San  Jose, 

Costa  Rica 


Abstract. — The  koinobiont  ectoparasitoid  Zatypota  petronae  Gauld  (Ichneumonidae)  parasitizes 
medium-sized  immatures  of  the  cobweb  spider  Theridion  evexum  Keyserling  (Theridiidae).  Zatypota 
petronae  apparently  attacks  the  spider  inside  its  retreat.  An  egg  is  glued  on  the  antero-lateral  dorsal 
section  of  the  spider's  abdomen.  First-instar  larvae  remain  partially  inside  the  egg  chorion  which  is 
attached  to  the  spider's  abdomen.  In  later  instars,  a  layer  of  a  brownish  material  (saddle),  to  which 
the  7th  and  8th  abdominal  segments  of  the  larva  adhere  ventrally,  anchors  the  larva  to  the  spider.  In 
the  last  instar  the  saddle  includes  the  egg  chorion  and  the  shed  exoskeletons  of  previous  instars.  A 
row  of  retractile,  dorsal  protuberances,  crowned  with  hooklets,  is  present  on  abdominal  segments  1 
to  8  of  the  final-instar  larva.  The  larva  uses  the  hooklets  to  grab  silk  lines  of  the  retreat  of  the 
spider's  web.  Hanging  on  the  spider's  web  the  larva  kills  the  spider  and  sucks  out  its  body  tissues. 
Then  the  larva  pushes  vigorously  laterally  with  its  head  against  the  spider's  corpse,  and  alternately 
presses  the  corpse  against  the  saddle.  These  movements,  in  combination  with  peristaltic 
movements,  free  the  larva  from  the  saddle  that  falls  to  the  ground  with  the  dead  spider.  The 
larva  then  constructs  its  pupal  cocoon.  Prior  to  cocoon  construction,  the  larva  induces  the  spider  to 
reinforce  the  retreat  by  adding  more  threads.  Parasitism  rate  and  host  behavior  are  also  described. 


The  polysphinctine  pimplines  are  koino- 
biont ectoparasitoids  of  spiders  in  several 
families  (Nielsen  1923,  1932,  Fincke  et  al. 
1990,  Hanson  and  Gauld  1995,  Gauld  et  al. 
1998).  Nielsen  (1923,  1932)  described  in 
detail  the  behavior  of  the  larvae  and  hosts 
of  several  European  polysphinctine  spe- 
cies. The  parasitism  rates  and  life  cycle  of 
Hymenoepiiuecis  robertsae  Gauld  on  the 
neotropical  tetragnathid  Nephila  clavipes 
(L.)  was  described  by  Fincke  et  al.  (1990). 
However,  larval  behavior  of  neotropical 
polysphinctine  wasps  has  been  described 
in  detail  for  only  one  species,  H.  argyr- 
aphaga  Gauld  on  the  tetragnatid  Plesiometa 
argyra  (Walker)  (Eberhard  2000a,  2000b, 
2001). 


Current   address:    Department   of   Entomology,    Kansas 
State  University,  Manhattan,  Kansas,  USA 
'  Author  for  correspondence 


Parasitoid  wasps  of  the  cosmopolitan 
speciose  polysphinctine  genus  Zatypota 
Forster  parasitize  spiders  in  at  least  five 
families  (Dictynidae,  Agelenidae,  Tetra- 
gnathidae,  Araneidae  and  Theridiidae) 
(Shaw  1994,  Gauld  et  al.  1998).  In  the 
neotropics  the  only  two  host  records  were 
Theridion  species:  T.  contreras  Levi  for  an 
unidentified  Zatypota  species  (Jimenez 
1987)  and  T.  evexum  Keyserling  for  Z. 
petronae  Gauld  (Barrantes  and  Weng  in 
press). 

The  larval  behavior  of  Zatypota  sp. 
(Jimenez  1987)  differs  in  some  aspects  from 
that  of  European  polysphinctine  species 
(Nielsen  1923,  1932)  and  H.  argyrapmaga 
(Eberhard  2000a).  The  larva  of  Zatypota  sp. 
was  said  to  hold  on  to  the  spider  by  biting 
the  dorsum  or  sides  of  the  anterior  section 
of  the  spider's  abdomen.  This  description 
is  likely  wrong  as  detailed  descriptions  of 
the  behavior  of  the   larva   of  Z.   albicoxa 


Volume  16,  Number  2,  2007 


327 


(Nielsen  1923)  and  the  larva  of  H.  argyr- 
aphaga  (Eberhard  2000a)  show  that  some 
posterior  segments  of  the  larvae  lodge 
ventrally  in  a  "saddle",  probably  coagulat- 
ed spider's  hemolymph,  that  adheres  tight- 
ly to  the  spider's  abdomen  (Nielsen  1923, 
Eberhard  2000a).  There  is  no  further  in- 
formation on  the  biology  of  larvae  of  this 
Zatypota  species.  Here  we  describe  the 
intensity  of  parasitism  and  behavior  of 
the  larva  of  Z.  petronae  and  its  host  T. 
evexum.  We  describe  for  the  first  time  how 
a  polysphinctine  larva  frees  itself  from  the 
spider's  corpse. 

MATERIALS  AND  METHODS 

Field  observations  were  made  from 
October  2005  to  October  2006  in  a  250  m2 
plot  in  the  understory  of  a  middle-eleva- 
tion wet  forest  patch  (9  54'N,  84  03' W; 
elevation  1200  m),  the  Reserva  Biologica 
Leonel  Oviedo  on  the  Universidad  de 
Costa  Rica  campus,  San  Jose  Province, 
Costa  Rica.  All  spiders  (or  nearly  so)  from 
third-forth  instar  outside  the  egg  sac  to 
adults  were  checked  for  parasites  every 
two  weeks;  most  spiderlings  disperse  from 
the  mother's  web  at  fourth  instar.  The 
small  eggs  and  early  instar  larvae  probably 
went  undetected. 

Theridion  evexum  constructs  most  webs 
between  0.20  to  1.5  m  above  the  ground 
(Barrantes  and  Weng  in  press),  making  it 
possible  to  find  practically  all  webs.  Addi- 
tionally, we  collected  seven  parasitized 
spiders  and  kept  them  on  their  webs 
indoors  to  observe  the  behavior  of  larvae 
and  spiders.  In  two  cases  we  transplanted 
the  plant  on  which  the  parasitized  spider 
had  constructed  its  web  indoors,  allowing 
us  to  observe  the  larva  and  host  behavior 
with  little  disturbance.  The  complete  larval 
development  was  not  observed  in  all  cases, 
so  sample  sizes  are  not  always  the  same. 
Behavior  and  morphological  features  of 
more  than  10  larvae  were  observed  under 
a  dissecting  microscope.  Video  recordings 
of  behavior  were  made  using  a  Sony  DCR  - 
VX    1000    camcorder    with    +5    close-up 


lenses.  Drawings  of  larval  behavior  were 
traced  from  video  recordings.  Voucher 
specimens  of  wasps  and  spiders  were 
deposited  in  the  Museo  de  Zoologia  of 
the  Universidad  de  Costa  Rica.  Wasp 
species  names  follow  Gauld  et  al.  (1998). 

RESULTS 

Percentage  of  parasitism. — Only  juveniles 
of  T.  evexum  were  found  to  be  parasitized 
by  Z.  petronae.  A  second  instar  larva  was 
feeding  on  a  juvenile  spider,  possiblv 
a  third  instar.  However,  final  instar  larvae 
were  found  feeding  on  large  immature 
spiders,  possibly  juveniles  of  fourth  to  fifth 
instars.  The  parasitism  in  T.  evexum  was 
very  low  (mean  percentage  of  parasitism/ 
census  =  1.39%,  SD  =  1.80,  n  =  53  bi- 
weekly censuses).  The  reproduction  in  T. 
evexum  is  extremely  seasonal,  and  the 
abundance  of  immature  spiders  (4th  instar 
or  larger)  susceptible  to  attack  by  Z. 
petronae  increased  in  March  and  declined 
drastically  through  August  (Fig.  1).  Be- 
tween September  and  February  the  popu- 
lation consists,  first,  of  mature  females,  and 
then  of  very  small  spiderlings  (Fig.  1). 
Parasitized  spiders  occurred  primarily 
from  March  through  August. 

Spider  web  and  wasp  attack. — The  web  of 
T.  evexum  includes  a  folded  leaf  that  forms 
a  conical  retreat,  with  a  tangle  in  front  of 
the  retreat  opening,  and  long  viscid 
threads  extending  from  the  tangle  to  other 
leaves  (Barrantes  and  Weng  in  press).  An 
additional  tangle  is  constructed  by  the 
spider  inside  on  the  upper  side  of  the 
retreat. 

We  witnessed  one  attack  bv  a  female  Z. 
petronae  wasp.  The  wasp  approached  the 
web  and  hovered  in  front  of  the  spider's 
retreat  opening.  The  wasp  then  flew  inside 
the  retreat.  A  few  seconds  later,  the  spider, 
with  the  wasp  perched  on  its  dorsum, 
dropped  about  10  cm  below  the  retreat, 
and  hung  on  its  dragline.  They  struggled 
for  a  few  seconds  and  then  the  wasp  flew 
out  of  sight.  The  spider  began  to  climb 
towards   the  retreat  but  after  advancing 


328 


Journal  of  Hymenoptera  Research 


350  n 


Dec    Feb    Apr     Jun    Aug    Sep    Nov 

Date 

Fig.  1.     Temporal  changes  in  the  number  of  immature  spiders  (black  rhombus),  males  (open  circles),  and 
reproductive  females  (black  circles)  of  T.  evexutn. 


about  four  centimeters,  it  became  para- 
lyzed and  fell  back,  motionless  for  about 
10  min,  dangling  from  its  dragline.  The 
spider  recovered  its  motion  slowly,  and 
with  clumsy  movements  cleaned  some  of 
its  legs  before  ascending  to  the  retreat. 
When  we  returned,  30  min  later,  the  spider 
had  fully  recovered  its  mobility.  We  did 
not  ascertain  whether  the  spider  had  an 
egg  on  its  abdomen. 

Larvae. — We  observed  one  egg  of  Z. 
petronae  glued  on  the  antero-lateral  dorsal 
section  of  the  abdomen  of  a  spider  collect- 
ed in  the  field,  a  first  instar  larva  emerged 
about  four  hours  later.  All  eleven  larvae  of 
different  instars  checked  under  the  dissect- 
ing microscope  were  attached  by  their  rear 
end  to  the  cuticle  of  the  antero-lateral 
surface  of  the  spider's  abdomen  (Fig.  2A). 

The  first  instar  larva  (n  =  3)  had  its 
posterior  end  lodged  inside  the  egg  chori- 
on, with  its  head,  thorax,  and  some 
abdominal  segments  protruding;  the  cho- 
rion remained  attached  to  the  spider.  In  the 


"second"  instar,  larvae  (n  =  5)  were 
completely  outside  the  collapsed,  flattened 
egg  chorion  that  was  embedded  in  an 
apparently  rigid,  semitransparent  layer  of 
brownish  material  (Fig.  2B)  (the  "saddle" 
of  Nielsen  1923).  The  ventral  surface  of  two 
or  three  posterior  abdominal  segments 
rested  on  the  saddle.  In  subsequent  instars, 
the  cuticles  of  the  previous  molts  became 
incorporated  into  the  saddle  as  they  ad- 
hered to  its  upper  surface,  against  the 
ventral  surface  of  the  larva.  The  egg 
chorion  was  near  the  spider's  surface,  but 
not  in  contact  with  it.  The  saddle  was 
attached  by  a  short  pedicel  to  the  spider's 
abdomen  (Fig.  2B),  and  the  larva's  abdom- 
inal segments  7  and  8  secured  it  to  the 
saddle.  Feeding  scars  were  observed  on  the 
nearby  dorsal  and  lateral  surface  of  the 
spider's  abdomen  (Fig.  2B). 

In  the  final  instar,  larvae  had  dorsal, 
two-lobed,  retractable  tubercles  on  eight 
abdominal  segments  (1st  to  8th);  these 
structures  were  absent  in  previous  stages. 


Volume  16,  Number  2,  2007 


329 


A 


* 


larva 


B 


y<i 

V 


■shed  cuticles 
spider's  abdomegj 


feeding 
scars 


Fig.  2.  Larva  of  Z.  petronae:  A-  Second  instar  larva  attached  to  the  antero-lateral  surface  of  the  spider's 
abdomen.  B-  Penultimate  larva  with  the  saddle  attached  to  the  spider's  abdomen.  The  shed  cuticles  of  previous 
molts  are  visible  under  the  larva.  Feeding  scars  (black  dots)  are  also  visible  on  the  surface  of  the  spider's 
abdomen.  (Photo  of  a  specimen  in  alcohol). 


When  extended,  the  tubercles  were 
crowned  with  a  circle  of  tiny  hooks  that 
allowed  the  larva  to  grab  the  threads  of  the 
spider's  web  inside  the  retreat.  The  larva 
could  extend  or  retract  independently  each 
lobe  of  the  tubercle,  and  the  tubercles  could 
be  retracted  rapidly  and  completely  into 
a  pocket.  Based  on  size  and  morphology, 
we  discerned  three  instars  in  the  larvae  of 
this  wasp.  However,  the  saddle  of  what  we 
thought  was  a  second  instar  larva  included 
the  chorion  and  the  shed  cuticles  of  two 
molts.  Hence,  further  observations  are 
needed  to  confirm  the  number  of  instars. 

The  final  instar  larva  spent  about  18  h 
attached  to  the  spider  (n  =  2),  three  to  six 
hours  after  removing  the  saddle  and  prior 
to  cocoon  construction  (n  =  5),  and  nearly 
18  h  constructing  the  cocoon  (n  =  1).  The 
duration  of  the  larva  inside  the  cocoon 
before  pupation  was  not  recorded.  One 
penultimate  instar  larva  molted  during  the 
night  and  the  next  morning  hung  from 
lines  near  the  roof  of  the  retreat  with  its 
dorsal  hooks,  and  fed  on  the  spider  for 
about  eight  hours.  During  approximately 
the  first  four  hours  the  spider's  legs  moved 
slightly,  but  later  we  could  not  detect  any 


movement.  The  larva  fed  first  on  the 
spider's  abdomen,  then  on  its  cephalotho- 
rax.  When  discarded,  the  spider's  carcass 
was  nearly  completely  empty;  even  its  legs 
were  almost  transparent.  The  larva  was 
thus  capable  of  extracting  nearly  complete- 
ly the  spider's  internal  tissues,  presumably 
using  capillarity  (Eberhard  et  al.  2006). 

Dislodging  the  saddle. — After  the  larva 
had  finished  feeding,  it  began  to  free  itself 
from  the  saddle  while  hanging  inside  the 
spider's  retreat.  The  process,  which  lasted 
about  two  hours,  included  three  types  of 
movement:  pressing  the  spider  carcass 
against  the  saddle,  pushing  the  spider 
carcass  laterally,  and  peristaltic  move- 
ments of  the  larva's  abdominal  segments. 
The  pressing  and  peristaltic  movements 
seemed  to  be  more  frequent  and  intense 
just  before  the  spider  carcass  and  saddle 
were  completely  removed. 

Pressing  movements:  The  ventral  side 
of  the  larva's  head  pushed  on  the  spider's 
anterior  end,  steadily  pressing  the  spider's 
carcass  against  the  saddle  until  it  bent 
almost  completely  over  the  saddle 
(Fig.  3A,  B).  The  larva  then  released  the 
pressure  completely  as  it  moved  its  head  to 


330 


Journal  of  Hymenoptera  Research 


Fig.  3.  Movements  of  the  spider  to  free  itself  from  the  saddle  (traced  from  video  images).  Pressing  movement: 
the  larva  places  its  head  near  the  spider's  chelicerae  (A)  and  presses  the  spider  carcass  against  the  saddle  (B). 
Lateral  pushing:  the  larva  places  its  head  on  the  anterior  tip  of  the  spider  carcass  and  pushes  it  laterally  (C).  A 
backward  final  push  completely  dislodges  the  saddle  from  the  larva  (D);  the  grey  arrow  shows  the  position  of 
the  larva's  head  tip  before  pushing  the  saddle.  Dotted  and  dashed  lines  represent  the  initial  positions  of  the 
larva  and  the  spider's  carcass  respectively. 


the  initial  position,  and  then  either  made 
another  pressing  movement  or  pushed  the 
spider's  carcass  laterally  (see  below). 

Lateral  pushing:  The  larva  bent  ventral- 
ly  until  the  lateral  section  of  its  head 
contacted  the  legs  and /or  cephalothorax 
of  the  dead  spider,  and  then  pushed 
laterally  (Fig.  3C).  Then  it  moved  its  head 
back  slightly,  maintaining  contact  with  the 
carcass,  and  pushed  laterally  again.  The 
larva  pushed  repeatedly  up  to  10  times 
before  reorienting  its  head;  the  complete 
carcass  moved  visibly  with  each  push  by 
the  larva.  The  larva  often  placed  its  head 
on  the  opposite  side  of  the  spider  during 
successive  pushing  bouts.  During  the  last 
three  pushing  bouts  the  larva's  head  was 


oriented  at  about  30  to  its  longitudinal 
axis  and  contacted  the  saddle,  rather  than 
the  spider  and  the  force  exerted  by  the 
pushing  movement  was  toward  the  rear  of 
the  larva's  body  rather  than  laterally 
(Fig.  3D). 

Peristaltic  movements:  Peristaltic  waves 
moved  posteriorly  along  the  larva's  abdom- 
inal segments  during  pressing  and  pushing 
movements.  The  last  segment  stretched 
extensively  backward  as  the  wave  reached 
it.  The  peristaltic  waves  were  strongest 
during  the  last  pressing  and  lateral  move- 
ments of  the  larva. 

Final  events:  As  soon  as  the  saddle  was 
released  the  larva  rubbed  its  head  against 
the  ventral  surface  of  the  segments  that 


Volume  16,  Number  2,  2007  332 

had  been  connected  to  the  saddle,  which  All  57  cocoons  found  were  constructed 
were  covered  with  a  mucilaginous  sub-  inside  the  spiders'  retreats,  but  their 
stance.  The  small  processes  on  the  ventral  attachment  varied  among  retreats:  71% 
larval  segments  that  are  inserted  in  the  were  attached  to  the  threads  of  the  tangle 
saddle  in  other  polysphinctines  (the  "taps"  near  the  retreat's  roof,  20%  were  attached 
of  Nielsen  1923,  Eberhard  2000a)  were  not  to  the  threads  applied  by  the  spider  at  the 
visible  in  the  Z.  petronae  larva  at  the  apex  of  the  leaf-cone  (Fig.  5A),  and  9% 
moment  the  saddle  was  released.  Howev-  were  in  the  middle  of  the  retreat,  attached 
er,  examination  of  two  saddles  under  the  to  a  thick  silk  cable  formed  by  several 
dissecting  and  compound  microscopes  independent  threads  (Fig.  5B). 
showed  a  wedge-like  depression  inside  Enemies  of  the  wasp. — Of  the  57  cocoons 
the  saddle.  This  depression  was  likely  found,  we  observed  two  predation  attacks 
produced  by  an  abdominal  projection  that  and  a  possible  parasitoid  attack  on  a  third 
anchored  the  larva  to  the  saddle.  cocoon.  One  pupa  was  attacked  by  Sole- 
Cocoon  construction. — One  larva  of  Z.  nopsis  ants  inside  the  spider's  retreat.  A 
petronae  began  cocoon  construction  at  second  pupa  or  larva  inside  its  cocoon  was 
about  18:30  h  inside  the  spider's  retreat,  attacked  by  a  penultimate  male  of  T. 
after  resting  for  nearly  two  hours.  We  did  evexnm  that  fed  on  the  immature  wasp 
not  follow  cocoon  construction  in  detail,  through  the  cocoon  silk.  The  third  cocoon 
but  our  incomplete  observations  indicate  had  a  lateral  hole  near  its  bottom  that 
that  the  behavior  was  quite  similar  to  suggested  the  exit  of  a  parasitoid,  as  adults 
cocoon  construction  by  H.  argyraphaga  of  Z.  petronae  exit  the  cocoon  by  cutting 
(Eberhard  2000a),  except  that  no  suspen-  a  circular  slit  near  the  cocoon's  upper  end. 
sion  line  was  built.  Construction  lasted  Host  spider  behaviour. — The  spiders  ear- 
nearly  18  h  (N  =  l).  It  began  with  the  larva  rying  first  and  possibly  young  second 
hooked  by  its  dorsal  tubercles  to  the  silk  instar  larvae  (N=4)  were  capable  of  cap- 
threads  of  the  tangle  inside  the  retreat  turing  prey  trapped  on  the  long  viscid  lines 
(Fig.  4A).  of  their  webs.  Their  attack  behavior  was 
The  larva  built  the  cocoon  by  attaching  indistinguishable  from  the  attacks  of  non- 
a  silk  line  (or  lines)  produced  from  its  head  parasitized  spiders  (Barrantes  and  Eber- 
to  the  tangle  of  threads  made  by  the  spider,  hard  in  prep.).  However,  spiders  with 
and  pulling  its  head  from  this  point  to  the  a  large  penultimate  instar  or  a  final  instar 
next  attaching  point,  which  was  either  larva  did  not  attack  prey  that  adhered  to 
another  tangle  thread  or  one  of  its  own  the  sticky  threads.  The  stickiness  of  the  last 
previously  produced  lines.  Cocoon  con-  capture  threads  produced  by  a  spider  with 
struction  began  around  the  posterior  por-  a  large  penultimate  instar  larva  was  nota- 
tion of  the  larva  (Fig.  4B)  and  then  gradu-  bly  reduced,  as  Drosophila  flies  (with  their 
ally  extended  upward  until  it  enclosed  the  wings  cut)  walked  easily  along  these 
larva.  The  first  silk  lines  around  the  larva  threads. 

formed  a  loose,  fluffy  mass  (Fig.  4C),  but  On    four   occasions   we   observed    that 

after  some  hours  a  much  denser  wall  began  when  a  larva  apparently  bit  the  cuticle  of 

to  form  around  the  larva  (Fig.  4D).  The  a  spider's  abdomen,  the  spider  jerked  and 

larva  frequently  paused  during  the  con-  tried  unsuccessfully  to  reach  the  larva  with 

struction  for  up  to  2  min.  After  20  h  the  its  legs  I,  II  and  III.  This  suggests  that  the 

larva   ejected   its   meconium   through   the  spider  perceived  and  was  irritated  by  the 

circular  hole  at  the  bottom  of  the  cocoon,  wounds  produced  by  the  larva.  In  one  case 

The  recently  constructed  cocoon  had  a  pale-  the  spider's   leg   II    touched   the  anterior 

yellow  color  that  turned  to  orange-yellow  portion  of  the  larva,  and  the  larva  imme- 

over  the  next  day.  diately  moved  its  anterior  portion  toward 


332 


Journal  of  Hymenoptera  Research 


Fig.  4.  Sequence  of  cocoon  construction.  A)  Final  instar  larva  recently  freed  from  the  saddle,  dorsal  tubercles 
are  visible  on  two  abdominal  segments.  B)  Larva  about  45  min  after  cocoon  construction  began.  C)  Cocoon 
construction  after  approximately  2  h.  D)  Cocoon  after  20  h;  note  the  meconium  below  the  cocoon. 


the  dorsal-middle  section  of  the  spider's 
abdomen  (out  of  range  of  the  leg)  and 
apparently  bit  her  again.  Examination  with 
a  hand  lens  showed  that  there  was  a  tiny- 
shiny  spot,  presumably  of  hemolymph, 
where  the  larva  had  apparently  first 
bitten  the  spider  (documenting  that  the 
larva  actually  bit  the  host  rather  than 
just  touched  it  with  its  mouthparts  is  not 
easy). 

The  web  retreats  housing  cocoons  had 
additional,  non-sticky  thick  threads  either 


across  the  retreat  opening  (72%,  n  =  57; 
Fig.  5B),  inside,  more  or  less  in  the  middle 
of  the  retreat  (20%)  (Fig.  5B),  or  both  (8%) 
across  the  retreat  opening  and  inside  it 
(Fig.  5B).  In  one  case  the  threads  inside  the 
retreat  were  so  dense  that  they  formed 
a  sketchy  sheet  just  below  a  cocoon 
(Fig.  5C),  which  was  attached  to  the  tangle 
threads.  A  parasitized  spider  added  more 
threads  to  the  apex  of  the  retreat  (Fig.  5A), 
possibly  during  the  last  two  nights,  before 
being  killed  by  the  larva. 


Volume  16,  Number  2,  2007 


333 


Fig.  5.  Retreat  constructed  by  Theridion  evexum.  A)  The  arrow  shows  the  threads  that  maintain  the  leaf-retreat 
folded.  B)  Threads  added  by  the  spider  at  the  retreat  opening  (a)  and  in  the  middle  of  the  retreat  (b).  C)  Sketchy 
sheet  in  the  middle  of  the  retreat.  Larva  of  Zatypota  petronae  induces  T.  evexum  to  produce  threads  at  the  retreat 
opening,  inside  it  and  to  increase  number  of  threads  that  maintained  the  leaf  folded  (black  arrow  in  A). 


DISCUSSION 

The  morphology  and  behavior  of  the 
larva  of  Z.  petronae  are  quite  similar  to 
those  of  larvae  of  other  polysphinctine 
species  (Nielsen  1923,  1932,  Fincke  1990, 
Gauld  et  al.  1998,  Eberhard  2000a).  How- 
ever, they  often  differ  in  where  and 
possibly  how  their  cocoons  are  attached 
to  the  host  web.  The  larva  of  Z.  petronae 
attaches  its  cocoon,  which  lacks  a  suspen- 
sion line,  to  silk  threads  inside  the  retreat 
of  T.  evexum,  Hymenoepimecis  spp.  attach 
their  cocoon  to  the  spider  web  (e.g.  N. 
clavipes  and  P.  argyra)  with  a  suspension 
line  (Fincke  et  al.*1990,  Eberhard  2000a, 


2001),  whereas  the  larva  of  Reclinervellus 
nielseni  (Roman)  [=  Polysphincta  nielseni] 
(Nielsen  1923,  Gauld  and  Dubois  2006)  and 
P.  gutfreundi  Gauld  (Gauld  et  al.  1998), 
which  also  lack  suspension  lines,  attach  the 
cocoons  to  the  threads  near,  or  on  the  hub 
of  the  orbicular  web  of  Cyclosa  conica 
(Pallas)  (Nielsen  1923)  and  Allocyclosa 
bifurca  (McCook),  respectively.  These  dif- 
ferences are  likely  determined  by  the 
characteristics  of  the  web  of  each  spider 
species,  particularly  by  the  modifications 
of  the  web  (the  "cocoon  web"  of  Eberhard 
2001)  induced  by  the  parasitoids  (e.g.  T. 
evexum  re-enforcing  its  retreat). 


334 


Journal  of  Hymenoptera  Research 


There  are  also  differences  in  how  larvae 
adhere  to  the  saddle.  Larvae  of  Z.  petronae 
apparently  adhere  to  the  saddle  using 
wedge-like  projections  of  one  or  two  seg- 
ments, rather  than  taps  as  in  Z.  albicoxa  and 
H.  argyraphaga.  Differences  may  also  exist  in 
the  sensitivity  of  the  host  to  the  wounds 
caused  by  the  parasitoid.  For  example,  P. 
argyra  did  not  show  any  reactions  to 
apparent  bites  of  H.  argyraphaga  larvae 
(Eberhard  2000a).  However,  T.  evexum 
reacted  by  jerking  its  body  and  moving  its 
legs  toward  the  point  where  the  larva  was 
biting  the  spider's  cuticle.  This  suggests  that 
chemical  composition  of  secretions  could 
vary  among  parasitoid  species.  Further 
research  to  confirm  chemical  differences  in 
the  saliva  of  parasitoids  and  differences  in 
sensitivity  of  spider  hosts  to  the  bites  of 
their  parasitoids  is  needed. 

The  release  of  the  saddle  by  final  instar 
larvae  is  much  more  complex  than  simply 
the  muscular  movements  of  the  posterior 
end  of  the  larva  as  suggested  by  Nielsen 
(1923)  and  Eberhard  (2000).  Without  the 
powerful  pressing  and  pushing  move- 
ments of  the  larva  against  the  saddle,  the 
peristaltic  abdominal  movements  are  pos- 
sibly insufficient  to  free  it  from  the  saddle. 
More  information  is  needed  to  examine  the 
possible  differences  among  polysphinctine 
species. 

The  larva  of  Z.  petronae  induces  the  host 
spider  to  add  more  threads  on  different 
sections  of  the  retreat  (apex,  inside,  and 
across  the  retreat  opening)  that  make  this 
structure  stronger  and  more  durable.  Add- 
ing threads  near  the  apex  of  the  retreat  is 
apparently  a  repetition  of  a  subroutine 
used  in  the  construction  of  the  retreat  by 
an  unparasitized  spider,  since  threads 
applied  in  similar  fashion  allows  the  spider 
to  fold  the  leaf  and  maintain  the  retreat's 
shape.  Similarly,  threads  across  the  retreat 
opening  were  occasionally  present  (3  out  of 
17  webs)  when  pre-adult  female  spiders 
were  molting,  though  these  threads  were 
not  as  abundant  as  those  in  retreats  of 
parasitized   spiders.   However,   the   thick 


cable  of  silk  threads  produced  inside  the 
retreat  was  not  found  in  webs  of  un- 
parasitized spiders.  The  reinforcement  of 
the  retreat  with  additional  silk  threads 
possibly  increases  the  protection  of  the 
cocoon,  primarily  against  heavy  rains, 
which  is  likely  important  for  the  wasp's 
survival.  If  a  retreat  opens  up,  it  is  unlikely 
that  the  thin  threads  of  the  tangle  inside  the 
retreat,  where  most  cocoons  were  attached, 
could  survive  heavy  rains  intact. 

Our  observations  suggest  that  Z.  petronae 
is  not  specialized  on  a  particular  species  of 
host.  This  wasp  parasitized  intermediate 
sized  spiders  (at  least  4th  instar),  but  the 
reproduction  of  T.  evexum  is  highly  sea- 
sonal and  large  juvenile  spiders  occur  only 
during  five  or  six  months  of  the  year.  Thus, 
it  is  likely  that  Z.  petronae  must  parasitize  at 
least  one  other  species  of  spider  to  main- 
tain its  population. 

The  percentage  of  parasitism  of  T. 
evexum  (1.39%  ±  1.80)  was  relatively  low 
when  compared  with  other  spider  species. 
Fincke  et  al.  (1990)  reported  that  the  annual 
percentage  of  parasitism  for  intermediate- 
sized  juvenile  females  of  N.  clavipes  was 
15-30%,  and  Eberhard  (2000)  reported  that 
the  parasitism  on  P.  argyra  was  higher  than 
40%  for  mature  females  and  higher  than 
3%  for  mature  males.  The  low  parasitism 
on  T.  evexum  also  suggests  that  Z.  petronae 
is  possibly  using  other  spiders  as  hosts  in 
the  same  area. 

ACKNOWLEDGEMENTS 

We  thank  Ingi  Agnarsson  and  Paul  Hanson  for 
identifying  the  spider  and  wasp  respectively,  and 
William  G.  Eberhard,  Paul  Hanson,  Mark  Shaw,  and 
an  anonymous  reviewer  for  valuable  comments  on 
previous  drafts.  We  also  thank  Andrea  Bernecker  for 
her  comments  that  greatly  improved  the  drawings. 

LITERATURE  CITED 

Agnarsson,  I.  2004.  Morphological  phylogeny  of 
cobweb  spiders  and  their  relatives  (Araneae, 
Araneoidea,  Theridiidae).  Zoological  Journal  of  the 
Unman  Society  141:  447-626. 

Barrantes,  G.  and  J.  L.  Weng.  In  press.  Natural  history, 
courtship,   feeding  behaviour  and   parasites  of 


Volume  16,  Number  2,  2007 


335 


Theridion  evexum  (Araneae:  Theridiidae).  Bulletin 
of  the  British  Arachnological  Society. 

Eberhard,  W.  G.  2000a.  The  natural  history  and 
behavior  of  Hymenoepimecis  argyraphaga  (Hyme- 
noptera:  Ichneumonidae)  a  parasitoid  of  Plesio- 
meta  cirgyra  (Araneae:  Tetragnathidae).  journal  of 
Hymenoptera  Research  9:  220-240. 

.  2000b.  Spider  web  manipulation  by  a  wasp 

larva.  Nature  406:  255-256. 

.  2001.  Under  the  influence:  webs  and  building 

behaviour  of  Plesiometa  argi/ra  (Araneae,  Tetra- 
gnathidae) when  parasitized  by  Hymenoepimecis 
argyrophaga    (Hymenoptera,    Ichneumonidae). 
journal  of  Araclmologi/  29:  354-366. 
-,   G.   Barrantes,   and   J.    L.   Weng.   2006.   The 


mystery  of  how  spiders  extract  food  without 
masticating  prey.  Bulletin  of  the  British  Arachnolo- 
gical Society  13:  372-376. 
Fincke,  O.  M,  L.  Higgins,  and  E.  Rojas.  1990. 
Parasitsm  of  Nephila  clavipes  (Araneae:  Tetra- 
gnathidae) by  an  ichneumonid  (Hymenoptera, 
Polysphinctini)  in  Panama.  Journal  of  Araclmologi/ 
18:  321-329. 


Gauld,  I.  D.  and  J.  Dubois.  2006.  Phylogeny  of  the 
Polysphincta  group  of  genera  (Hymenoptera: 
Ichneumonidae;  Pimplinae):  a  taxonomic  revision 
of  spider  ectoparasitoids.  Systematic  Entomology 
31:  529-564. 

,  J.  A.  Ugalde  G.,  and  P.  Hanson.  1998.  Guia  de 

los  Pimplinae  de  Costa  Rica  (Hymenoptera: 
Ichneumonidae).  Revista  Biologia  Tropical  46 
(Supl.  1):  1-189. 

Hanson,  P.  and  I.  D.  Gauld.  1995.  The  Hymenoptera  of 
Costa  Rica.  Oxford  University  Press,  Oxford. 

Jimenez,  M.  L.  1987.  Relaciones  entre  arahas  y  avispas. 
Folia  Entomologica  Mexicana  73:  173-183. 

Nielsen,  E.  1923.  Contributions  to  the  life  history  of 
the  pimpline  spider  parasites  (Polysphincta,  Za- 
glyptus,  Tromatobia).  Entomologiske  Meddelelser  14: 
137-205. 

.  1932.  The  Biology  of  Spiders.  Levin  &  Munks- 

gaard,  Copenhagen. 

Shaw,  M.  R.  1994.  Parasitoid  host  ranges.  Pp.  111-144 
in:  B.  A.  Hawkins,  and  W.  Sheehan,  eds.  Parasitoid 
community  ecology.  Oxford  University  Press,  Ox- 
ford. 


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